Download Feeding, ranging and social organisation of the Guinea baboon

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Feeding, Ranging and Social
Martin
Organisation
of the Guinea Baboon
John Sharman
F,
Thesis
submitted
a'
to the Department of Psychology,
University
St.
of
Andrews
SUMMARY
Before
almost
this
field
ninteen-month
feeding,
The troops
troops
other
with
other
in
the
compositions
both
they
troops
did
in
the
the
both
of
were
reports
carried
out
a
in
and aspects
Both
and
by comparison
large
their
although
was
troops,
study
age-sex
indication
some
that
was low.
troops
in
time
more
related
to
since no rain
fell
probably
dry
the
of
members
and
similar,
were
and feeding-in
spent
which were large,
productivity,
there
moving
when they
thesis
were recorded.
and
genus,
was
papio,
on foot,
to be unusually
population
more time
wet,
papio
possible.
exceptional,
adult
differences
These
in
budgets
spent
found
were
not
the
The activity
whenever
censused
troops
into
of
behaviour
and social
area,
were
P.
The monkeys were followed
ranging
were
This
environment.
of two troops
study
known
recruitment
natural
Senegal.
south-eastern
of their
its
in
unknown
baboon, Papio
C he Guinea or Western
study,
than
season
behaviour.
social
in
changes
seasonal
in six months of
the
year.
The home range of'one
square kilometres,
of the study
differences
troop,
roughly
dry
season
great
This distance
of baboons,
troops
most other
the
was
8 kilometres.
square
daily
and
large
frequently
more
than'they
through
those that
they visited
they
areas
they
visited
in
did
less
which
areas
with
that
frequently.
there
little
was greater
to
In
did others,
was relatively
shade.
In
dry
the
visited
moved
more
than through'those
season
dense
mean or
both
troops
shade morex, frequently
wet season
by
in
productivity
The troops
frequently
the
low
and
no
were
travelled
than that
in the wet.
habitats
the
about
variation
well,
by
day
either
per
travelled
and was ascribed
troops
There
kilometres.
the mean distance
in
there
although
18 to 20
about
covered
50
to
less
home
was
range
whose
the other'troop,
while
known, ranged over about
seasonal
troops
45
they
avoided
some
slowly
that
visited
than
areas
a
in which visibility
Sleeping
ranging
was poor.
were
sites
patterns
inversely
to
site.
afforded
them protection
restricted
to places
The baboons
baboon
troops
wet
known
when
The social
It
sexual
during
in
the
this
except
those
that
of
species
time
times
with
of. the
stress.
lived
Instead.,
the
female's
which
of
they
baboon.
partners,
other,
to mate with
constrained
species,
at
the
of
was unlikely
were
two other
for
beneath
and mating
organisation
with
compared
females
found
sites
were
resembled
the
in
and the
but
a
of
number
diet
in. their
Animals
boulders,
the
More than
was available.
study.
the
vertebrates
some
eaten.
also
gelada.
throughout
invertebrates
were mostly
were
types
which
throughout
changed
were known to be eaten,
increased
trees
they
this
diet
of foods
variety
items
food
different
food
a wider
in
sleep
nearest
and was more diverse
seeded,
being
water.
In
Their
the
sleeping
was permanent
areas.
fruited-or
plants
The
frugivorous.
in other
to
chose
predation.
there
which
from
distance
the
on
home ranges
the
of
usage
the
of
influence
profound
apparently
largely
studied
season,
hundred
in
a
with
square
against
were
as various
year,
the
baboons
The
have
to
baboons,
the
of
proportional
sleeping
found
is
of
system
baboons,
in
only
there
of
oestrus.
Adult
to be
consortships
baboons
males
with
which
adult
females
as are
appeared
formation
uncommon in
in
one male,
were
Theropithecus
including
society
a
baboons
these
in
competition
between
groomed
competitive
adults
each other
mating,
DECLARATION
I alone
been
submitted
higher
degree.
This
the
the
in
of
is
Doctor
General
No.
reports
or entirely
submitted
of
and as a candidate
Ordinance
it
thesis,
part
disseration
Degree
student
this
wrote
to
in
the
Philosophy.
for
12 in
my own work
I
September
of
St.
was admitted
the Degree of Doctor
1976.
has
it
application
any previous
University
and
not
for
Andrews
a
for
as a research
of Philosophy
under
CERTIFICATE
I hereby certify
of
research
under
ordinance
of
resolution
submit
of
under
that
supervision
General
the
No.
after
thesis
being
admitted
He has fulfilled
12.
Unversity
the accompanying
John Sharman has completed
Martin
Court,
in
1967,
application
as a research
the
for
the
Philosophy.
A. Whiten
Research
August,
1981.
Supervisor.
Degree
student
of
the
qualified
to
conditions
No. '1 and is
terms
nine
of
Doctor
Contents
Introduction
1.1
Chapter
2.1
Materials
2: Site,
Geography
2.1
2.1
2.2
2.3
2.4
Geology, Topography and Vegetation
Climate
Habitat Map
2.5
Estimating
2.6
Visibility
2.7
2.8
2.9
Cover
Topography
the
Following
Baboons
2.10
Nomenclature:
Age-Sex Classes
Chapter
1:
and Methods
3: Demography
Introduction
Group Size:
Group Size: Methods
3.3
Group
3.4
The Social
3.5
Age-sex
Structure
3.6
Age-Sex
Structure
3.7
3.8
Age-Sex Structure
Age-Sex Structure
3.9
Predation
3.14
3.15
Chapter
4.1
4.2
4.3
4.4
Chapter
5.1
5.2
5.3
5.4
5.5
5.6
2.7
2.10*
2.12
2.14
Chapter
3.1
3.2
3.10
3.11
3.12
3.13
2.6
Composition
Habitat
Size:
2.1
2.3
2.5
2.15
3.1
3.1
3.2
3.2
Results
Group at Mt.
Discussion
Assirik:
of
Troops:
of
Troops:
3.5
Introduction
3.9
Method
3.9
3.11
3.19
of Troops:
of Troops:
Results
Discussion
of Group:
Introduction
3.20
Predation and Size of Group: Results
Predation and Size of Group: Discussion
Food and Size of Group: Introduction
Food and Size of Group: Results
3.22
3.23
3.25
3.27
and Size
Food and Size of Group: Discussion
in
Changes
Causes
General Discussion
of
of
4: Activity
Budgets
Ecological
Constraints
on Activity
Activity
Methods
Budgets:
Activity
Results
Budgets:
Period:
Social
Activity
Budget within
Discussion
Size
4.1
4.1
4.2
4.4
4.9
5.1
5.1
5.1
5: Ranging
Introduction
Size of Home Range
5.2
Movements of Baboons
Day Range Maps
Size of Home Range:
Movements of Troops
Group
3.28
3.29
5.6
5.10
5.11
Results
Home Range Area
Movements of Troops
Habitat Variables
5.25
5.26
5.28
Chapter
6.1
6.2
6: Sleeping
Sites and Water
Introduction
Sites:
Sleeping
Results
Sites:
Sleeping
6.1
6.1
6.2
6.3
6.4
6.5
6.6
Sleeping Sites: Discussion
Water: Introduction
Water: Methods
Water: Results
6.7
Water:
5.7
5.8
5.9
6.5
6.5
6.8
6.9
6.13
Discussion
1
Chapter
7.1
7.2
7: Feeding
Work
Previous
Omnivory and Terrestriality
7.3
7.4
Comparison of "Direct
Boulder-Rolling
7.5
Methods:
7.6
7.7
7.8
Damage to Plants
Comparison of Diets
Omnivory
Results:
7.1
7.1
7.2
Observation"
with
"Feeding
Boulder-Rolling
Damage to Plants
Comparison of Diets
7.12
Discussion:
7.13
7.14
7.15
7.16
Breadth and Nature of Diet
Boulder-Rolling
Damage to Plants
Overlap in Diet between Baboons
Chapter
8.1
8.2
8.3
8.4
8: The Influence
of Food on Ranging and Troop
Influence
of Food Species on Ranging
Methods
Results
Discussion
Chapter
9.1
9.2
9.3
9: Social
Organisation
Review of Earlier
Work
Foraging
Groups
Associations
between Age-Sex
9.4
Intertroop
Encounters
Chapter
10: Mating
Systems
10.1
Review of Earlier
Social
7.7
7.7
7.8
of Baboons and Chimpanzees
Biasses
of
Baboons and Chimpanzees
in
the
7.22
7.25
7.25
7.27
Record
Feeding
7.28
7.31
7.32
7.34
and Chimpanzees
Size
8.1
8.1
8.2
8.5
8.8
9.1
9.1
9.5
9.11
Classes
9.19
10.1
10.1
Work
10.4
Groups
10.3
10.4
10.5
10.6
10.7
Nearest Neighbours
Inter-Individual
Distances
Male and Female Interactions
Grooming
Summary
Chapter
11: Ecology,
Social
7.4
7.4
7.5
of Data
Collection
7.9
7.10
7.11
10.2
Remains"
10.8
10.10
10.13
10.15
10.21
and Mating
Organisation
P. papio
2
System of
11.1
The colour
his
his
is
forms
cheeks
middle
the
and hands
face
of his
surface
of
sized
are
black,
and his
tuft
a considerable
body is
dog;
his
by no means dull
(Papio
common baboon
but
upper
on
each
are
thickset
From:
Bennett
proportions
baboons,
abandoned
all
the
and
In bulk
he is
under
to a
equal
but
and inelegant;
of
he
or inactive.
the mighty
with
The hair
side;
E. T. The Tower Menagerie,
(quoted
... amongst
reddish-brown;
white.
eyelids
covered.
sparingly
is
papio)
their
pretence
granite
almost
at
crags
and boulders
human eyes
finesse.
fixed
"Run for
Hill,
was a vast
on the
itl"
in
1970).
assembly
invaders.
of
Biggles
he yelled.
From: Johns W.E. (1936)
Biggles
in Africa.
ACKNOWLEDGEMENTS
With
I
pleasure
great
who have helped
many people
for
preparation
Whiten,
field,
the
home during
their
valuable
who have
baboons
this
me into
am deeply
their
grateful
shared
with
of
Jezebel,
small
indebted.
and
life
me in
with
who taught
of
pursuit
Hall,
Land Rovers
who later
Alexander
and
a
who shared
I am even
baboons.
cylinder
six
those
Byron
Stephanie
to
I
was
settlement
all
of
Baldwin
Pamela
isolated
this
For
of West Africa
corner
companionship
grateful
me more about
100 kilometers
Our camp was about
Simenti,
where
Patrice
Marty
thanks
for
their
to
grateful
than
so many
in tending
Lucazeau
National
to
Andre
Parks,
they
On my return
the
I
I thought
to
am grateful
and
to
for
staffs
Senegal
Shirley
Strum
evenings.
Without
have been a great
my warmest
them,
deal
and
drier.
invaluable
and
advice
to
and to
work
Park and of
that
in
the
the
the
Director
of
Niolcolo-Koba
for
Hotel
Simienti
sites
in Kenya.
work possible.
I visited
for
I extend
their
To
so good.
Orbell
Senegal,
of
of the
Joe Popp and Jeanne
tasted
never
of
permission
in making
road
Hotel
from
vehicle.
Government
Dupuy,
provided
from
gave
would
freely
of an ageing
Park and to the
the help
baboons,
also
the ills
I am grateful
refreshing
"en brousse"
life
pool,
Gazelles
and Nigel
Fernandes
spirtually
dusty
and
hot
of
and colder
and Michel
swimming
National
Storks
cold
They and Claude
all
in
Life
have met the
to know.
wanted
help
in a remote
me
help.
his
few years.
and
Mike and Sarah Harrison,
I am especially
her
they
me into
given
for
never
last
the
which
with
ease
community
us, especially
Anderson.
year
the
by them and by the
made enjoyable
it
for
and
opportunity
accepted
Jim
occupied
for
so fully
my life
my
when I
welcomed
grateful
I would
Tutin
McGrew and Caroline
in
Senegal
Andy has also
and I am very
work,
in
kindly
to civilization.
the
of
encouragement
and
Whiten
all
thesis.
this
my enthusiam
He and Suzie
of my written
Bill
Without
gave
my readjustment
critisism
me help
and rekindled
had begun to get discouraged.
thanking
of
opportunity
I was preparing
me while
Andy
My supervisor,
this
take
Altmann
inviting
baboon
three
for
introducing
me to
spend
me to
several
I
their
weeks
to know the Pumphouse Gang.
getting
This
me into
thesis
his
extremely
the
in
unit
I benefited
College
Dunbar
me to
baboons,
which
Reasearch
Group disbanded
from
dedicated
the
Albon
and Glen
them,
and
and
Iason
P.
Way.
help
the
members of
of
to
to
permission
Large
I recieved
that
by Tim,
I
there,
Steve
up.
To
extend
my
of my writing
work
other
his
join
generated
bulk
the
the
When King's
me to
The encouragement
I am
Wrangham.
Richard
thesis.
this
invited
me through
for
which
relation
atmosphere
enthusiastic
Horne
for
from
in
papio
permeates
Storey's
boosted
Gabriel
to
see
Tim Clutton-Brock
Group at
the
welcomed
Anatomy,
and especially
now
perspective
from
greatly
Group,
Research
encouraged
Mammal Research
of Veterinary
Department
the
grateful.
King's
Robin
was written
Chivers
where David
up in Cambridge,
thanks.
It
is
with
encouragement
will
be able
ever
friendship
during
Mackinder
with
has
Snelson
given
on
commented
thesis
Hawksworth
least
emotional
the
for
the
his
of
my life.
of
this
in
and
Duncan
this
read
and
Debbie
Lee has
Phyllis
and has read,
thesis.
both
thesis,
thoroughness.
advice,
I
that
hospitality
references.
and practical
chapter
I doubt
Rasmussen has kindly
painstaking
and
assistance,
Cambridge.
preparation
of checking
every
help,
years
Kathy
with
support
in
sociable
in
chapters
the tedium
To all
reread,
these
grateful.
The University
This
Gareth
photography.
on and critisised
the resources
found
enormously
several
the
acknowledge
I have
the
of
and with
I am deeply
people
repay
one
undertook
me both
to
helped
computing
commented
friends
the
of
I
that
pleasure
Cambridge
of
to carry
Computing
Service
in chapters
out much of the analysis
and
typeset
was funded
initially
was edited
using
gave me the means and
programs
and five.
three
by Philip
developed
Hazel.
The research
Great
like
grant
I am grateful
Britain.
to
thank
ended.
my mother,
It
is
for
their
by the
generous
who has supported
to her that
I dedicate
Science
Research
support.
me openhandedly
this
thesis.
Council
Finally,
since
of
I would
the
SRC
CHAPTER 1: Introduction
Papio
little
papio
is
known
the
contrast,
the closely
The
taxonomy
priority
of
(Linnaeus
1758),
1758),
like
Theropithecus,
is
are
(1975)
in
is
restricted
having
took
Most
lies
place
previous
P. hamadryas
single
are
adult
thought
groups
been
in
subgroups
male.
to
their
(1975b),
a small
in
Senegal.
in
In
1766),
(Erxleben
Papio
1777).
and other
(1977),
Delson
and
of
in
these
case
be an adaptation
unproductive
and Guinea
Guinea
The National
has
the
arid
mating
the
of
to
studied
in
central
semi-desert
foraging
in
this
in
is
(Kummer
in
Ethiopia.
of
restricted
these
within
to
organisation
dispersed
widely
1968a).
and
several
groups
social
small,
Eastern
east
large
females
P. hamadryas
habitats
1.1
of
in
been
live
species
this
which
of the species.
have been conducted
1835)
but
Bissau,
in
Park
found
being
of West Africa,
distribution
which
the
the
(Linnaeus
Mandrillus
of the distribution
(Rüpell
studied
Both
(Linnaeus
Delson
area
Mali,
restricted
highlands.
Ethiopian
which
has
Delson
thesis
P. hamadryas
Theropithecus,
Papio,
take
treat
and
and
about
this
species,
to
of
of baboons
studies
(1970)
genus
history
in the centre
its
The
1960)
should
Throughout
P. cynocephalus
closely
Mauritania,
T. gelada
within
sites
of
1827),
the genus Papio
five
serious
debate
considerable
and Hill
of
from
Tappen
1963,
1970).
up
made
over
range
Africa.
Southern
is
its
of
are among the best
(1954).
distributed
areas
most
study
revised
of
related
in
and Patterson
P. papio
forms
P. papio.
and evolutionary
is
there
in Hill
(Lesson
and
cercopithecids
of
species
Apart
Freedman
(1967)
was
1792),
The taxonomy
Andrews
it
P. anubis
(Kerr
P. ursinus
1970),
and Napier
though
as
living
1843),
clear.
not
1956a,
the recognised
Napier
Papio
genus
Hill
is
1966, and review
follow
shall
(Booth
1975a,
(Buettner-Janusch
baboons
the
level
taxonomic
(Geffroy
By
primates.
nomenclature
(Delson
I
of
and
captivity.
outside
the genus, and the single
genus Theropithecus
non-human
of
behaviour
or
ecology
species
other
problems
which
of
related
known of all
its
in the wild,
been studied
1820) has seldom
(Desmarest
The
extinct
a
ýý
ulw
Mauritania
Senegal
Dakar
.
......
.......... ...
........
....... Tamýbýacoýrada
....
..!
0
...
Mali
.
ASSIRIK; ...
-""""o
If N
ssou
............
Guinea
Figure
^
ý_
\
1.1(a):
distribution
of
io a io;
(Mt
Assirik)
location of field site
Gem
Western
A'ýergarthecus gelada
or
Guinea
........:.:......:.:.
.......:.........................
4,0
ý:.
...:::..:
Olive
%
00
Yellow
i
cynocWwAa
'.
or
000
01
::::::
""""""
Figure
ohaºyas
Chocma
1.1(b):
distribution
baboons
long-tailed
the
approximate
of
after July and &"n
1973
T. oswaldi
lived
and
in
The other
(Fig
1.1).
desert
in
Kenya.
and southern
in
from
to arise
thought
females.
To succeed
kin
These
groups
certain
females,
males
in the troop.
wooded
grassy
but
of
in
societies
similar
it
lives
advanced
identical
studies
Plio-pleistoscene
grass
baboons
P. papio.
baboons
edges
of
not
Our
knowledge
riparian
1981),
but
studied,
of
baboons
the
1.2
of
in
sites
possible
lives
it
The
Although
desert,
woodland
in which
of
is
groups
one-male
much
and
1980b).
most
tree-studded
of
reconstructions
covers
coastal
in semidesert
P. hamadryas.
scrub
which
in Namibia,
study
autonomous
other
lightly
in
(Anderson
an environment
range
with
by
Tanzania
these
suggest
which
protection
similar
and desert
forests,
kin.
with
contested
In one locality
for
other
with
has been studied
have been in
of
(McGrew et al.
been
and
in
of
into
proposed
environment
swards
have
Papio
that
food
mutual
the
of P. hamadryas
those
fragmentation
to
on the
or
grasslands,
of
In all
of P. anubis.
resembling
this
in
semi desert
swamp.
to those
for
found
P. ursinus
is
relationships
special
often
Kenya
trees,
the Okavangu
in groups
previous
are
for
are
consortships
Southern
for
coalitions
groups
form
may
males
scattered
on the edges
effectively
other
has been studied.
with
kin
organisations
environments
plains
reason
with
in
they
to
who tend
organisation
social
form
in groups
live
to
females,
to compete
competition
sexual
social
P. cynocephalus
this
Adult
predators.
Identical
in
of central
wooded grassland
This
females
in Uganda, and
has been done on this
receptive
relatives.
in Ethiopia,
forest
been found
sexually
the need for
associate
against
with
female
close
with
associate
species
has always
for
distributions
wooded gorges
in Kenya, riparian
place
compete
males
adult
which
This
thinly
in the thinly
has taken
however,
The one-
more extensive
Most of the work that
in Tanzania.
forest
species,
forest
was
organisation
social
adaptation.
far
in
has been studied
does
as
range,
of low productivity.
of this
have
baboons
Cameroon, coastal
a lakeside
This
1981).
be
a relic
may
well-known
P. anubis
own
to environments
an adaptation
of T. gelada
male groups
(Dunbar
today
home
have
to
thought
are
species
its
with
each
groups
patas
also
probably
other
perhaps
one-male
Erythrocebus
Theropithecus
the
interspersed
present-day
the
adaptations
range
of
of
the
social
has
rights
been
the
promiscuous
with
one-male
This
aspects
of
the
troop
is
population
in
animals
are
patterns
of
of habitat,
diet
one of
social
For
chapter
by
use,
sake
is prefaced
their
the
two baboon
species
of P. papio,
several
apart
in
turn
vegetation,
factors
of p. papio
of
by the ecology
to
clues
thought
clarity
by a detailed
no further
influence
to influence
the
organisation.
I
within,
review
by various
is
Since
and water.
social
organisation
Finally
at length.
this
given
the
which
influences
was influenced
sleeping
system
to
extent
of
in which
of the area
sites
of
budgets
activity
social
ranging
introduction.
1.3
the demography
Environment
is discussed
and mating
the
of
nature
and
number
The
in
organisation
social
the
some detail.
the way in which
of,
influence
environment.
which
the major
organisation
to
may provide
including
the diet
the
with
the members of the group,
influenced
are
examined
aspects
the
from
set
influences
in
examined
and hence
range
in primates
mating
species
in one population
thought
between
affected
have therefore
is
it
exclusive
the
would
and place
structure
a group
lives
are
which
relationships
group
This
to examine,
out
sets
ecology
individuals
1975).
have
groups.
Since
possible
been limited.
P. papio
of
males
baboons
savanna
study
baboons.
(Boese
various
this
to
and
environments,
has therefore
that
suggested
females
over
to
in particular,
environment
It
the
of
organisation
important
baboons
I examine
one population.
here,
but
each
Figure
2.1:
in
field
lateritic
area.
plateaux
Extent of
Plateau,
North
the
of
southern'rim
Camp was on
to
Plateau
South
its
approach
closest
near
Hidden
valley
Plateau
Mt Assirik
CHAPTER2: Site,
Materials
and Methods
INTRODUCTION
2.1: Geography
is
Senegal
by Mauritania,
Bissau.
the
the
in
an area
of
the park
flat
whose
Assirik
and Guinea
lies
park
to
top,
in the
savanna
south
highest
in
the
the
about
0.5
sq km, is
only
311m above
represent
the
north-western
surroundings
source.
The area
in which
km, with
Mt. Assirik
lying
massif,
this
point
in which
took
study
1981).
of its
lies
Most of
Mt. Assirik,
Mt.
level.
sea
limit
the
of
its
Gambia finds
occupied
place
It
in the north
park,
River
the
and to the east
within
and Guinea
sq km.
(McGrew et al.
of
the Fouta-Djalon
of
197,000
Sudan savanna
with
north
(GPN du NK), 500km from
Senegal's
woodland,
on the
by Guinea
south
seaward
of
its
and
foothills
8000 sq km (4%) of
about
bounded
du Niokolo-Koba
National
drought-deciduous
of
and on the
by Mali,
east
Pare
The Grand
sea, covers
the
western-most
on the
in Africa,
country
100 sq
roughly
centre.
2.2: Geology, Topography and Vegetation
A5 to 10m thick
chemical
surface
tropical
weathering,
was overlain
in the middle
a broad ribbon
of ancient
this
In the field
ribbon
area this
lay
plain
a plain
(Fig 2.1)
plateaux
to the south, overlying
to
stretching
To
1971).
and Michel
the
Atlantic
by an Upper Cambrian
was underlain
of plateaux
lay an area of low
of Middle Cambrian pelites.
Almost
communities
no soil
that
had formed on the
they supported
Andropogon pseudopricus).
with
laterite
rock (Hebrard
To the east of the ribbon
red sandstone.
hills
raised
metamorphic
This
This low mesa was
boulders.
isolated
of
product
summit of Mt. Assirik.
formed the flat
of similar
ferrugineous
in a 10km-wide band from the north
which stretched
Ocean.
a reddish,
by a cover of residual
of a series
the west of
of laterite,
crust
stunted
were generally
In places
bushes, principally
Boscia
senegalensis
called
"Combretum
and other
scrub"
or
laterite
"scrub
2.1
plant
grasses
shallow-rooted
(eg
these grass swards were interspersed
Combretum etessei,
species
and the
plateaux,
(Adam 1971).
on
laterite
but
This
plateau".
including
also
I
have
Debris
had
habitat
accumulated
boulder
as
Many of
plateaux.
abyssinica)
africana,
these
communities
I
Bridges
These
1970).
generally
more or
Combretum
spp.,
Pterocarpus
Hexalobus
included
In
lowest
the
horizons,
Plant
but
there
in
the
of
whose trees
dry
Water draining
basins
catchment
in
rested
This
was the
and
abyssinica),
of
only
Permanent
plateaux.
of
were
these
ravines,
or
permanent
dense
obligate
riverine
in
water
the
drinkers,
gallery
2.2
area,
but
forest,
larger
of
forest
dry
spp.
incised,
into
with
five
of
one
the
through
cut
water
typically
30 m wide
flowed
deep
and
from rim to rim).
and was not
with
the
Along
(Andropogon
near-permanent
also
Grewia
scattered
flowed
were
gullies
sandy soil.
ravines
which
in places.
and Zizyphus
in the
of Mt. Assirik
outlets
layer
deeply
erosion
active
of
illuvial
lacked
grass
Oncoba spinosa,
the
of
consisting
strips
and
of
season.
west
(Adam 1971).
were characteristically
from the sides
other
with
elephant
of
areas
were
Grewia
(Adam 1971).
the
from
arrecta
as much as 30 m deep and only
only
to
areas
included
wet
the
organic
distant
aethiopum,
riverbeds
bottom
(sometimes
primates
strips
the
areas
superficial
of Pobeguinea
there
whose
plains
as "scrub-grassland"
Borassus
laterite
surrounding
narrow
most
it
banks of up to 2m showing
vertical
or
porous,
(Oxytenanthera
included
The seasonally
highly
tufts
riverbeds
bamboo
gayanus),
and
plains
the
to
deeper,
I described
among the
2m in
and Hyparrehenia
In
wooded,
Many species
the
in
(Adam 1971,
lightly
madiensis.
about
the
variably-
thonningii,
and in
particles.
the
less
basins
was a substantial
made up a habitat
sides
was
loose,
The community
to
reaching
Schizachyrium,
sand-sized
communities
lasiodiscus
trees,
the
of
soil
unconsolidated
and Vitex
beneath
debris
Piliostigma
erinaceous,
Andropogon,
the
plateaux
about
these
parts
3m and 4m tall.
monopetalus,
grew beneath
Genera
were
and
woodland".
both
of
consisting
areas
touched,
canopy
to the formation,
or granitic
senegalensis,
often
"closed
or
schistic,
most
grasses
forest"
quartzitic,
between
lasiodiscus,
Khaya
Canopies
a lithosol
of
trees
with
digitata).
the
of
edges
by bamboo (Oxytenanthera
cordifolia,
and had led
was active
unconsolidated,
Cola
"dry
called
and elsewhere,
slopes
sized,
(eg
trees
the
at
cliffs
stabilised
were
and Adansonia
on slopes
talus
slopes
large
and
Afzelia
Erosion
slopes
these
the
beneath
only
nourished
Ceiba
vital
to
restricted
pentandra,
Erythrophloem
Adansonia
mespiliformis,
Saba
(Pan troglodytes
chimpanzees
area
which
They also
prep).
with
papio)
as sleeping
The geology
in
is
taken
the
rock
gives
of various
sizes
the
diet
iron
of
(Henty
aethiops
and were
by baboons
used
the
of
in
sabaeus)
vegetation
to
rock
the
environment,
split,
vertebrates
such
rock
these
boulders
swells.
Repeated
forming
the
are
rich
In the dry
season
and drying
boulders
of
boulders
provided
for
and
and snakes,
many
of
part
a major
what was possibly
over
is
the
season
layer
These rough
(chapter
to
wet
wetting
a surface
as lizards
Baboons found
by rolling
In
and aluminium.
(Weyman and Weyman 1977).
small
surface
shrinks.
and
importance
of
and the
oxides
water
of invertebrates.
species
their
its
up
causes
for
patas
patas)
consequence
in
stable
oxides
up by these
eventually
shelter
in a field-
details
area had another
chemically
hydrated
partially
water
the
of
Laterite,
baboons.
1979)
prep),
Further
sites.
allowed
(1979).
in Baldwin
given
in
and several
forests
(Cercopithecus
monkeys
including
(Baldwin
(Erythrocebus
(Harrison
territories
productive
(Papio
green
provided
liana,
gallery
to survive
patas
Diospyros
heudelotti
These riverine
verus)
supported
also
Landolphia
(Adam 1971).
of Combretum
of
and many species
latifolia,
spp.,
Ficus
africana,
digitata,
Nauclea
senegalensis,
species
Afzelia
suaveolens,
7).
2.3: Climate
Adam (1971)
"The dry
indistinct
overheated
the
rains.
is
season
soil
The
torrents,
the
implacable
of
savanna
lift
that
like
2m tall,
the
grass,
the
are
now
everything
a
dust
animals
vault
used to obtain
area have been described
high
through
in detail
2.3
the
dry
the
forests
which
the exuberant
climatological
the
everything...
vanish,
mires,
beneath
soil
into
colourless,
skies,
black
with
ash,
atmosphere,
the
The vegetation
hiding
become
disappears
by fire,
a furnace....
grows
mudflats
sun, white
scorched
grass
baked
The methods
field
the
radiating
beneath
glitters...
season
whirlwinds
disappear
shadeless
the
at the site.
of the seasonality
description
a vivid
horizons,
trees,
skeletal
gives
data
by Baldwin
awakens with
dirt
the
the
had
and the climate
been
barely
sun
vegetation.
(1979).
become
gullies
which
roads
"
of
the
A summary and
Figure
2.28
.
Climate
at
(This
figure
230
Assirik
It,
is
not
referred
to
in
the
text)
rainfall
150
100
50
0
JFMAMJJASOND
'C
40
öaiy
mean
max
temperature
u
90
3o
80
is
20
daily
mean
min
JFMAMJJAS0ND
701
information
some additional
Air
temperature,
hourly
between
Annual
(November
in
fell
in
noon
April).
typically
humidity
dry
the
was highest
humidity
Relative
in
and low
September)
storms
the
month
(about
season
Most
occurred
at
(about
85%
wet season
15% at
dawn and lowest
at
year
The wettest
and most
in
to be high
tended
the
(198mm and 277mm).
twice
20-50mm,
of
assessed
study.
no rain.
with
330mm) and September
storms,
Relative
night.
at
once
(with
were dry,
to April)
were
6 months of
5 or
1000mm, but
was about
rainfall
the field
throughout
daily.
were collected
humidity
relative
and
07: 00 and 20: 00 hrs
or December
was July
rain
temperature,
sun
data
Rainfall
follows.
noon
in
shortly
after
around
15: 00-
noon.
Mean daily
16: 00hrs.
varied
mean daily
range
usually
between
1978)
of
shade
of the hottest
temperature
and coolest
of the day.
(1965).
1972, only
any other
The
calculated
large
given
site
the
means
in the
temperatures
from Thompson
data are taken
recorded
at two
1969,
Nathan
Dunbar
and
used by
and Waza in
the Cameroons (the
March had more hours of bright
sunlight
August had the fewest
mean annual
Mt.
evapotranspiration
from the annual rainfall
by Holdridge
(996 gm m2 yr
1)
site
1970) There were 6.6±1.56 hours of bright
month (an average of 9 per day) while
developed
productivity
line
and Gartlan
per day on average.
sunlight
(the
The mean daily
between
range
while
07: 00hrs,
around
in the dry season reached levels
Badi
-
1978),
1977).
100C above air
insolation
100km from Mt. Assirik),
used by Struhsaker
method
months was about
6.5°C.
(April
at
occurring
and the
The following
Sun- temperatures
41°C
and
was 11°±3.8°C
rose to roughly
baboon field-sites
other
usually
at
occurring
1977) and 27°C (April
20°C (November
Sun temperatures
heat
(August
29°C
temperature,
minimum
between
varied
maximum temperature,
at
was then
in Rosenzweig (1968).
The mean annual
estimated
The standard
(Rosenzweig 1968).
2.4
error
from
(4).
(640mm) was
Assirik
and mean annual temperature,
(1962).
than
using the
above-ground
the
on this
regression
estimate
is
2.4: Habitat
Map
with
within
km which
a rectangle
area,
(scale
area
they
types,
1: 50,000)
in
were viewed
they
stereo
a resolution
allowed
from
were
of
six
when
trees,
(1979)
Baldwin
than
finer
types
that
individual
of
this
of the
photos
detailed
showed some canopies
habitat
A map of
aerial
sufficiently
together
be enclosed
could
by 11 km east-west.
was prepared
which
be used,
known to
was not
10 km north-south
of
7 habitat
showing
field
and
15 sq
about
of baboons,
troops
The area known to be used by the two study
attempted.
The
habitats
correspondence
depicted
they
that
Correspondence
Appearance
on photo
is given
zones
in Table
on
the
photographs
and
the
2.1.
Pale grey
grey
Nearly
Habitat
type
grain
almost
little
Stands of grass, usually
pan
always on laterite
pale grey;
slight
Combretum scrub with grass;
pan
usually on laterite
pale;
little
Grassland
darker;
grainy
Scrub
pale;
grainy
Open woodland with
understory
dark;
little
Woodland.
Often with bamboo
Some areas closedunderstory.
canopy dry forest
white;
To draw the
map, acetate
The lines
corresponding
of
paper
transition
grassland
gallery
was placed
over
grass
forest
matched pairs
of
between zones of equal density
to each of the 7 habitats
2.5
but not
scrub or trees
with
with
Riverine
none used
photos.
Photographs
density
black
and texture
Between Zones on Aerial
Types
and Habitat
Appearance of
subdivisions
Very pale
aerial
the
2.1:
Table
Dark
between
were traced,
with
a
2.1:
Plate
Habitat
types
Scale roughly
in field
area.
1.3cma per km.
tý
%Apo
'äx.
KEY:
black
pale
dark
pale
dark
pale
dark
yellow
yellow
green
green
pink
pink
riverine
gallery
grass
on laterite
Combretum
scrub
open woodland
dry forest
scrub
scrub
grassland
t
't
wvF
".
.
ý'
forest
plateau
on laterite
plateau
.,
Rotring
onto
pen,
by overlaying
of
these
1 km.
5 cm to
(Plate
Each
was coloured
zone
the
the
and
in
to
result
result
a scale
photographed
The
sheets.
intersected
I
in
The aerial
the
photos
distance
the
near
on a line
the
of
those
When the
the
of
baboons
of
this
whose
was
error
distortion
edge of
the
both
which
of
another,
and which
photo.
Features
(n=40)
2.1±0.94%
by
estimated
was
features
of
terrain
the
of
appearance
and centre
a mean of
were
the
outstanding
local
the
lay
near
together
closer
centre.
less
but
lines,
were
errors
all
enlarged
probably
most
two
one photo
to
map was
was
error
position
that
the map
Since
1 mm.
accuracy,
this
of
effect
edge of
photo
near
distorted
between
perpendicular
edge
final
line
mapped
patterns
with
acetate
overlain
every
than
ranging
with
slightly
The
edges.
the
comparing
occurred
acetate
small.
satisfactorily
towards
of
less
determine
rarely
could
was made by
error
the
across
versions
was normally
association
this
of
drawn
were
two
the
line
a transect
position
lines
zones
between two zones was
the ground,
sometimes
the map, each on a separate
of
part
of
between
gap
be used
to
photos.
"transect"
Several
sheet.
the
An estimate
into
an area
separate
the border
Occasionally
on
a gradation
reflecting
two versions
constructing
how to
decide
to
easy
and texture.
sometimes
resolution
poor
than
maps and enlarging
was normally
of density
indistinct,
the
individual
map was made
2.1).
It
was
acetate.
of
sheets
separate
The final
than
may have
40m on
about
been
the
in
more
scaled
also
in
the
portions
of
ground
certain
The
up.
the map.
2.5:
In
imposed
on the
samples.
explained
and
the
calculate
map a grid
of
area
each
quadrat
The
dimension
above,
the
of
"grain"
squares
then
was
into
25
0.16
2.6
of
the
the
into
(40m
hectare
because
map.
various
oriented
divided
40m was chosen
size
by
occupied
kilometre
1 km square
Each
east-west.
quadrats,
to
order
Composition
Habitat
Estimating
I
it
and
north-south
25
x
four-hectare
40m)
"point"
represented,
recorded
I
habitats
the
as
single
type that
habitat
I considered
sample.
The habitat
in
of
terms
point
composition
of each point
part
then be described
quadrat
could
entire
map in
and the
samples,
The final
samples.
Table 2.2.
each 4 hectare
of
25 point
to occupy the greatest
terms
the mapped area
of
The range given here assumes an error
56,250
of
shown in
is
of +5%.
Table 2.2:
Habitat
Composition
Habitat
% of map
hectares
0.4611
6.2(121
39.7
Riverine Gallery Forest
Grassland (usu. on laterite)
Scrub Grassland
Combretum scrub (on laterite)
41.7
587.0
37.7521.0-
659.7- 729.1
1233.5-1363.3
694.4
1298.4
22.5(74)
1915.7
2031.7
1819.9-2011.5
1930.1-2133.3
27.3(46)
2461.1
2338.1-2584.2
21.235)
Open Woodland
Visibility
Visibility
primates,
is
fragmenting
Harding
coped
1976,
with
troops
Popp 1978).
fragmentation
Aldrich-Blake
and
and
levels
of
In
attempt
Saayman
visibility
to
foraging
strategies
a baboon
on the
be able
into
to
Dunbar
1972).
There
which
promotes
with
ground
see another
data
of
a tree
baboon
in
2.7
no
these
and
have
apparently
strategy
which
et
1971,
al.
Kummer 1968a,
the
habitats.
used
strategies.
to
distances
the
the
of
foraging
be
1970,
Altmann
quantification
170cm above
various
seldom
as a unit,
1972,
may later
I measured
visibility,
and in
been
either
which
Nathan
where
sites
(Aldrich-Blake
and
has
consequences
for
foraging
a
groups
of
baboons
sites
adopting
small
1968,
provide
field
strategies
field
In
(Altmann
parties
by
visibility
have
forage
apparently
At other
foraging
1979).
Leland
foraging
small
poor
Tayler
an
baboon
the
turn
in
may
and
in
part
strategies
(Struhsaker
good,
into
includes
Crook
foraging
structure
visibility
an important
may play
and
social
to
error
estimated
559.0
7.706)
14.47)
Dry Forest
Scrub
2.6:
Area
Field
of
ground
correlate
over
which
was likely
The
method
to
used
estimate
the
MacArthur
and MacArthur
silhouette
of
imaginary
p,
an
density
of
a silhouette
If
enough
the
probability
distance
at
vegetation
half
which
then
would
of
to
the
obscure
that
the
encountering
an
along
view
1m
cylinder
such
by
used
1 sq cm and length
any
me to
enabled
to
probability
that
were
does
not
by
the
by
pd
e-
the
also
similar
area
be given
p=
The
the
sectional
would
manner
a
cross
d metres
after
in
large
distances
these
(1961).
object
cylinder
then
contain
foliage
determine
area
a board
of
was obscured
be h where
-ph
e
= 0.5
or
p=
This
relationship
obscuring
vegetation
"baboon-sized"
squares
x 50cm chequerboard
At
sampled.
these
watched
while
chequered
points
edge of
the
directions
the
board
in
contact
none of
nearest
5m; distances
visibility
the
park
in
described
same points
From each point
an observer
with
see (a)
board
moved along
periodically
placing
this
habitat
sampled
were
later.
2.8
with
its
the
four
lower
the
the
each of
four
to the board was recorded
eyes at about
in
taken
of the squares
100m were estimated
over
the
to
60cm and (2)
(1)
fewer than half
over 200m to the nearest
was not
In
ground.
just
Distances
vertical,
each of
from the observer
the board.
grassland
the
with
metre when the observer
and (b)
for
turn,
zone to be
the
so that
the
observer,
the compass in
of
170cm above the ground could
Scrub
holding
assistant,
the distance
the nearest
seasons.
50cm
bottom.
the vegetation
were built
cairns
10cm x 10cm
a regular
provide
the
at
within
well
in subsequent
facing
surface
cardinal
a 10cm x 50cm margin
points
an
to
A
habitat.
each
13 white
pasting
cardboard
of
were selected
be used again
could
was constructed
with
Random points
by
in
a of
area
cross-sectional
space
of
metre
a 50cm x 60cm piece
onto
the
determine
cubic
per
target
-1/h
us to
enables
0.5
x in
to the
50m.
wet
from
season.
a road-strip
The
results
survey
of
In
the wet season two baboons on the
were unlikely
to be able
scrub-covered
plateau
In
or
higher.
or
collapsed,
the
view.
can clearly
per
dry
leaving
be seen in
the
while
in
in
the
grass
in
the
in
was trampled
the
view
in
wet
the
area
the
of
season
silhouette
2.4).
tall
with
were
in
grasses
habitats
were the
season they
off,
obscure
and Open Woodland habitats
on Laterite
dry
burned
to
(Table
two seasons
height
chest
down,
and
in
difference
season was
wet
trees
shrubs
the wet season because of the
the
the
but Combretum
grew to
many places
obscuring
seasonal
in
Grass
Visibility
which
grass
of habitat
in any habitat
fire-resistant
only
of
dense in
obscuring
season
The effect
As a result,
visually
2.3).
by grass,
the
metre
cubic
areas,
(Table
obscured
principally
to see each other
20m apart
more than
ground
the
these
least
vegetation.
Table 2.3:
over
Visibility
Habitats.
in Various
which observer
could see specified
I
iI
Season
Wet
Dry
note:
I
Half Visible
Height of Eyes
60cm
170cm
Habitat median range median range
Zi
13
GP
4
1-12
1-10
1 28.5
SP
13-44
15-20
30
1 2.5
OW
1- 63
0- 7
I----I---SG
1 8.5
FB
0-18
10
1-17
1 12
RG
1-23
13
3-34
Invisible
Height
60cm
=
Imedian range
17
J 31.5
4.5
9
1 11
3-11
21-49
2- 76
4-13
3-19
GP
SP
154
22-200
54.5
25-2001 88
1 49
32-300
14-125
51
29-1751 58
OW
1 21
4-90
SG
I-
FB
1 23.5
17
RG
KEY
GP
OW
FB
RG
in metres
Distance
amount of chequerboard.
-
7-57
5-14
to habitat
types:
grass on laterite
plateau
open woodland
dry forest,
sometimes with
forest
riverine
gallery
Scrub
grassland
was not
21.5
-
21
10
4-68 1 55.5
30
-1
7-51 1 45.5
5-28 1 16
SP scrub on laterite
SG scrub grassland
bamboo understory
sampled
2.9
in
the
wet
of Eyes
170cm
median range
8
35
11.5
11.5
4-15
20-60
1- 9
7-16
0-48
83
32-300
30-200
60
26-120
26-84
67.5
23-143
57.5
25-200
58.5
24
24-73
5-66
0-100
22-65
5-39
plateau
season.
Riverine
forest
the
was
habitat
only
sometimes worse in the dry season than
In
2.3).
the
with
undergrowth
improved
in
the
In
some cases
the wet season (Table
change in
visibility
density
the
by light
was
streambed
as
such
obstacles
off
the
of
limited
the
near
carrying
was
visibility
as growth was presumably
season by water
wet
which
was in
was little
there
seasons,
by water.
and not
fallen
habitat
this
it
in
trees.
2.4:
Table
Square
of Silhouette
of Habitat
Centimetres
Cubic Metre
per
Season
I
Dry
above Ground
I Height
above Ground
60cm
170cm
mean
{
60cm
170cm
on laterite
2310
1733
2022
1
128
127
128
Scrub on laterite
Open woodland
243
2773
231
2310
237
2542
1
1
141
330
136
322
139
326
815
587
693
533
754
555
1
1
295
990
330
693
313
842
Wet
Habitat
Grass
Height
Dry forest
Gallery
Riverine
2.7:
Cover
Kummer (1968a)
whenever
lower,
have
might
after
the
influenced
wet and dry
the
noted
of
the
seasons
percent
presence
09: 00.
In
of
50°C
relative
in
The same transects
assessment
P. hamadryas
topped
sometimes
some 10°C
that
reported
possible
temperatures
in
mean
of
environment
where
sun
shade
in
temperatures
were
habitats
different
The amount
baboons.
of
shade
was measured.
At every
apart
shade
shade
of
assessing
cover.
foliage,
where
the
habitats
used for
of
and
amounts
ranging
four
an
sat
down in
baboons
from
were used in
visibility
metre
along
grass,
the
vertically
transect
the
I
above the
ground.
In order
chequerboard
to estimate
the effectiveness
was marked with
beneath areas on the transects
of this
cover,
100 5cm x 5cm squares,
assessed
2.10
as having
a 50cm x 50cm
and placed
cover.
at noon
The number of
in full
squares
40% of
About
level,
the
scrub
unsurveyed
full
sunlight,
shade, and in penumbra was recorded.
field
(I
area
have included
than 5% ground
had less
grassland)
20% had more than 75% (Table
while
in
the
figure
this
above grass
cover
2.6).
Table 2.5:
Percent
Grass on laterite
plateau
on laterite
plateau
Open Woodland
between
(1967)
proportion
the
of
penumbra
every
1-9
0-7
22-50
3
26
-
-
-
-
84
96
assessed
as having
method,
65-93
88-99
cover
changed
like
that
differences,
seasonal
6-42
80
93
75-98
92-100
A more objective
little
Emlen
of
as was suggested
amounts of shade found beneath such "covered"
case
decreased
branches
while
in
the
in
lit
from
in
drought-deciduous
The proportion
in
points
with
the
the
sun
ground
bore
on
season
were
leaves
in
season
2.5),
although
there
and cast
the
plateaux
gallery
a direct
were
deep shadows.
median
percentage
was not
statistically
In
forests
the
many
penumbra,
principally
limits).
confidence
riverine
(Table
dry
the
as
season,
dry
and least
of the ground in deep shadow in each habitat
from data
statistically
Table
dry
to
wet
wet
of
season,
dense
in
open woodland.
seasons was calculated
shadow was not
by
binomial
shadow was densest
the
on
the
on plants
In
most branches
directly
(95%
foliage
case.
shadows
decrease
significant
2.6
every
season
in
increased
area of
whose
wet
The apparent
ground
shade
of greater
naked
Table
3
two seasons.
consequence
the
1-4
have shown greater
might
In
the
2.5
samples
seasons.
by the different
the
1-7
32.5
Dry Forest (some bamboo)
Riverine Gallery Forest
The
Dry Season
range
median %
3
Grassland
Scrub
Habitats
Wet Season
median %
range
Habitat
Scrub
in Various
Cover
of
in Tables
correlated
2.4).
2.11
2.5 and 2.6.
with
the
visibility
in both
The amount of
(compare
Table 2.6:
Percent
Habitat
Intensities
of Ground in Various
Assessed as having
Beneath Points
Habitats
in Various
Illumination
13.5
Penumbra
25
6-35
Shadow
57
24-76
range
median
range
Full Sun
7-45
8.5
7-39
27.5
19-49
54
12-67
15.5
3-24
50.5
19-79
Penumbra
12
2-29
34.5
6-81
Shadow
73
51-95
10
0-33
Woodland Full
Sun
Dry For. Full
Riverine
Dry Season
Wet Season
median
Plateau
of Sunlight
Cover
Sun
8-47
20.5
6-34
69.5
40-88
44.5
34-68
6.5
7
2-15
1-23
9
15
8-13
3-29
86.5
68-94
76
58-88
Penumbra
14
Shadow
Full Sun
Penumbra
Shadow
17-42
33.5
2-46
9.5
Table 2.7:
Percentage
of
the
Ground
Grass on laterite
Scrub on laterite
Open woodland
Dry forest
Riverine
Gallery
field
ridges,
features
occurred
sleeping
sites
and
in
and dry
baboons,
while
constructed
the
the
Forest
ranging
included
area
ravines,
I
1.4
1.6
2.6
35.6
70.7
1.7
1.7
23.6
58.4
83.0
Topography
The
the
Shadow
Percentage of Dense Shadow
Dry Season
Wet Season
Habitat
2.8:
in
of
the
the
relief
various
I
streams.
each
season
water,
others
were
map of
baboons.
recorded
in
quadrat
2.12
the
which
field
such
10
of
area.
their
were
chosen
chosen
on the
same criteria
two
field
or
area
more
as
cliffs,
topographic
Two features,
for
the
If
features
natural
relevance
which
was used
features
occurred
to
by which
to
in
plot
the
I
same quadrat
in
higher
hierarchy
a
common features
their
this
ordering
feature
and their
relatively
by my assessment of
was modified
the
Firstly,
than
rank
was
which
principles.
higher
Secondly,
when none of
The categories
quadrat.
two
had
to the baboons.
"undifferentiated"
topographic
by
generally
although
importance
by the
determined
features
rare
relatively
it
characterised
a quadrat
other
9 features
relative
rank
was described
as
in
the
appeared
in
given
are
Table
2.8.
Table
2.8:
Types
Topographic
% of
in
No. of Quadrats
Area
Field
Feature
Rank
Quadrats
1
2
Sleeping
Site
Dry Season Water
19
21
0.8
0.8
3
Ravine
30
1.2
4
5
6
Cliff
Ridge
Catchment
364
14.7
7
8
Drainage
Marigot
9
Plateau
10
2.8.1:
Topographic
Site:
trees
all
were
held
contained
Cliff:
the
8.5
1043
42.1
course
a water
for
were known to
baboons
which
one or more pools
containing
most or
all
of
sleep.
dry
the
season.
in which
These pools
in or downstream of the ravines.
a narrow
sites
211
in the text.
are given
in
persisted
were usually
Ravine:
14.7
2.1
Definitions
Dry Season Water:
water
364
52
Stream
of Features
Sleeping
5.7
9.3
Stream
Undifferentiated
Definitions
140
231
in
cleft
a laterite
in
ravines.
permanent
water
found
laterite
plateaux
30m high,
where
vegetation
at
the
the
base
All
in
ravines
held
dry
season.
the
all
streams,
the
sleeping
and nearly
Ravines
usually
forest.
gallery
riverine
Nearly
plateau.
often
laterite
of
these
2.13
ended
crust
cliffs
in
cliffs,
had
been
occasionally
eroded
was typically
away.
dense,
up to
The
often
large
having
senegalensis)
Ridges:
(eg
trees
Catchment Streams:
dendritic
between Mt. Assirik
Streams:
Senegal
to
more or less
of
Undifferentiated:
with
Following
that
they
reached
later
their
sampling
including
all-day
that
the
Senegal,
these
plateaux
to
mean
areas
extensive
were
the
of
none
containing
of
low-lying,
of
these
sleeping
in
site
in
the
topographic
other
flat
soil
sandy
lay
quadrats
the
in
Data
methods
which
described
occasionally
Kew Gardens.
in
in
detail
with
the
reference
to
other
later
for
African
to
and
relevant
whenever possible
members of the Stirling
S. Hall
were made
according
collected
were
food were collected
but
Observations
notes were made in a notebook
catalogued.
by other
that
time
to the
the morning
and
are
them from the
to stay with
evening.
and longhand
usually
(SAPP),
I tried
on foot.
destination
Samples of
identification,
Project
it
use
area.
left
they
chapters.
I
the Baboons
transcribed
several
Here
in
used
and
plateau".
vegetation.
10x40 binoculars,
through
the
between
plains
"creek",
or
watercourse.
small
Most
Baboons were followed
time
the
across
an area
west of the field
basins
plateaux.
was normally
poor
the catchment
laterite.
naked
This
covered
2.9:
in
a quadrat
features.
which drained
"backwater"
laterite
"bowal"
called
Saba
more major rivers.
any
across
vines
hills.
and other
meandering
meaning
to
refer
"watercourse
Plateau:
word
and
(eg
slopes.
and the fringing
streams
French
a
talus
gullies
and one of three
ravines
Marigot:
the
stabilising
senegalensis)
ran out from Mt. Assirik
ridges
Drainage
Khaya
Primate
sources,
and A. Whiten accompanied me on several
follows.
The vegetation
restricted
I was normally
within
them.
Before
and to
judge
either
startled
the view of the baboons to such an extent
their
flight
I became used to following
their
pace and to
adjust
distance
mine accordingly,
or lost
see
than by vision,
more by noise
them and caused them to flee,
2.14
I
could
as
as soon
I
contact
repeatedly
with
them
in the attempt
end
the
of
not to "push"
them.
Habituation
study.
hampered by the observation
there
sleeping
sites,
followed
from one sleeping
the
that
study
by which
area,
knowing
tail)
and Fable's
troop
(after
the
study
In
animals.
seen more than
the troop
same troop,
or
animals
If
once.
in which
it
I
did
change
identification
the
Both
compared the behaviour
not
and ecology
also been collected
on the behaviour
2.10:
Age-Sex Classes
Nomenclature:
Since
entirely
category"
individuals
on age-sex
were for
categorisation
and "age-sex
class"
in
this
2.15
is given
on
was
a known troop
I assumed
seen was the
that
secondly,
troop
In
firstly,
supposes,
these
reare
suppositions
in
the dry
season,
thesis
I have
this
two troops,
though data has
was unknown.
not known, I have depended
thesis.
are used to refer
one of the groups whose description
marks
of baboons whose troop
the most part
short
characteristic
certain.
of these
named
were
distinctive
and
of
without
an equally
This
to one or other
was probable,
troops
field
marks
same troop.
certain.
follow
and subsequently
troops,
the
characteristic
seen in
was previously
Where I refer
questionable.
detect
with
was ever
of
not
to
to the
a distinctive,
with
male with
an individual
it
a day's
These
male
tried
the baboons
using
were
end of
following.
an adult
a fragment
were
the
an adolescent
some cases
identifications
the
been
had
(after
Throughout
that
I
shared
or subtroops
days belonged
troops
two
came to
rarely
troop
that
also
were
the wet season at the end of
was not until
clear
troop
tail).
that
I
troops
on two successive
site
became
time
which
Anecdote's
the
it
recognition
Lsince
the
common towards
no way of knowing whether
was usually
it
became less
individual
and
and
conditions,
As a result,
same troop.
This
The terms
to animals
below.
"age-sex
belonging
to
Age-Sex
Code
Class
Male
adult
Categories
at Mt.
recognised
Assirik
Description
male
Mane and canines
fully
may be broken or worn).
Gives characteristic
(M)
young male
Mane developed,
fully
that
developed
pooled
Oestrus
Pregnant
but
a fully
of
not
adult
female
adolescent
females
classified
as oestrus
female
female
callosities.
female with
(Altmann
of
with
male.
Canines
This
for
swollen
sex skin.
in prefertile
this
under
abdomen noticed
in
late
was
of
that
Note
cycles
were
criterion.
pink to scarlet
In
well
often
class
as
analysis.
Taken as sign
1973b).
as large
quite
males
Adult
as mane of
Body not
adult
oestrus
Adult
as large
and sharp-looking.
with
pregnant
Massive appearance.
bass "wahoo".
male.
adult
(canines
developed
para-ischial
pregnancy
pregnancy
swelling
morning
(before
early
foraging).
Ventral
female with
ventral
Dorsal
infant
Adult
female carrying
(often
black)
infant
(often
brown) on
on ventrum.
female with
Adult
dorsal
dorsum.
infant
infant
female carrying
2.16
Age-Sex Categories
Code
Class
Female
female
Description
Nipples
of
unknown
sexual
button-like
status
subadult
male
not
status
subadult
female
to
adult
distinguish
frontally
exposed.
older
juvenile
active
young
Usually
juvenile
Small
Brown I
brown infant
Hair
(almost)
Trans I
transitional
Hair
black
infant
flanks.
but
class,
only
may be
visible
Too large
but lacking
Very hard class
juvenile
Larger,
sometimes.
chest
Active,
an infant
appear.
females,
brown buttons,
Gangly.
Sexual
size.
grass
to
like
Nipples
older
Young J
long
starting
than
female build.
Older J
adult
and
pink
be missed.
might
ventrally
more muscular
Subad F
Full
in
recorded;
Mane and canines
hard
when nulliparous,
when parous.
elongate
carried
Subad M
(cont)
at Mt. Assirik
recognised
when
to be
full
adult
to distinguish.
but not as gratuitously
as younger juvenile.
independent,
but
Play is rough.
often
unweaned.
and active.
completely
on crown and spine,
Face black.
2.17
brown. Skin black.
brown on
Age-Sex Categories
Code
Black
at Mt. Assirik
recognised
Class
I
black
(cont)
Description
infant
Hair
or mostly
completely
black.
Skin pink or
red.
Ventr
I
infant
carried
ventrally
Dorsal
I infant
carried
of un-
on the
riding
usually
dorsally
Unknown animal
Infant
ventrum
of
another
female
an adult
on the back of another
Infant
riding
often
an adult
animal
animal,
female.
Animal whose class
could
not be determined.
known class
Juvenile
Infant
Juvenile
of
Juvenile
which could
unknown class
accurately.
infant
Independent
unknown
of
class
be made out.
2.18
infant
not be classified
whose colouration
more
could
not
CHAPTER3: Demography
INTRODUCTION
3.1: Group Size:
Introduction
(1977)
Dupuy and Verschuren
de NK often
would
be
probably
largest
the
assertion.
Data
P. papio
in the park.
Group size
is
(1977)
is
in this
provided
1979, Cohen 1969,1971,1972),
and Altmann
1970 Chepko-Sade
et al.
ecological
for
1963a, Suzuki
1977, Hall
pressures
individuals
example,
in
but more likely
predation
at sleeping
Washburn
mingle
sites
and
interact.
3.1.2: Nomenclature:
I distinguish
was counted
but
The
individuals
out of sight
is
size
and
fission
(Altmann
1978, Dunbar
et al.
(Altmann
by
largely
are governed
may be less
likely
1977);
to
suffer
competition.
to form large
the
of
often
groups,
1970, Saayman 1971c,
groups
may not
may or
of
composition
age-sex
one
such
here.
recorded
in which
the
age or
sex of
few or
in which the age-sex
I was able
of the troop
to detect
the
(by movement of the foliage,
or
censuses.
A partial
observation
conditions
reduced the certainty
census
3.1
the
number of
none of
the
individuals
individuals
of many or all
class
was
of the
was determined.
count or census was known to be incomplete.
counts
of
migration
and fusion
and Altmann
The members
18 troops
and Taylor
feeding
(Altmann
their
1970, Taylor
aggregate
between "counts",
in some fragment
Frequently
groups
to support
Counts and Censuses
and "censuses",
recorded,
large
sometimes
1961).
at Mt. Assirik
assemblage
and Altmann
and waterholes
and DeVore
of which
all
known.
(Altmann
term
1979, Cheverud
to experience
Savanna baboon troops
death,
troops
these
of
sizes
1963a, Nash 1976),
1979),
(Altmann
on the
and long
and Olivier
1968, Hall
1974d, Furuya
no data
in the GPN
troops
primate
of birth,
and Altmann
and Dunbar
were true,
provide
chapter
by rates
influenced
this
unprovisioned
Dupuy and Verschuren
Unfortunately
If
more than 500 members.
include
of P papio
troops
that
claim
presence
noises,
of animals
that
on)
so
so
and
These I describe
was also
with
which were
recorded
the
as "partial"
when poor
which individuals
could
be classified.
3.2: Group Size:
Methods
Counts were made whenever the baboons were sighted
to this
not unique
52 of
more than
two
field
observers
Sabater
Pi (1968),
(1971),
Rowell
undulating
open,
primate
topography,
was the major
discusses
further
duration
count
of
the
sort
vegetation
counts
an
partial
frequency
constructed
indicators
additional
in
changes
is necessarily
troop
smaller
and 87 counted
Repeated
in
of known groups
wet
season
remained
In
season,
35)
Counts
wet
wet
while
of. both
season
those
used
were
as
a "median
that
Thus if
than the group size.
27,309 429 63,64,70
with
The
but smaller
is not clear,
median counts.
roughly
In the dry season no constancy
count
size.
showed that
at a time.
the
thicker
and have
animals,
Note
of
counts
Results
counts
the
to
the median count would have been 63.
presumably tend to give smaller
3.3: Group Size:
fewer
counts
between median count and group size
relationship
groups will
on 7 occasions,
possible
made in
median
of a group of 100 were seen repeatedly,
fragments
and
I have excluded
9 or
whose
than
size,
Repeated
analysis
contained
distributions
of
of group size
a troop
which
was sometimes
of
(1970)
group
Counts
Ransom
problem
Aldrich-Blake
size.
In this
results.
counts
it
troop
of
and
rather
which,
an open plateau.
crossed
Jones
the
visibility,
Assirik
Mt.
indication
gave variable
or
count"
as it
troop
gave
between
relationship
At
vegetation,
at Mt. Assirik.
difficulty
observation.
a complete
this
groups
dense
of
(1963),
mention
all
no
by either
Homewood (1976),
(1969)
that
reported
Maxim and Buettner-Janusch
in
a problem
seen at any one time
and Struhsaker
observing
(1962a)
Hall
country.
(1969),
Maples
(1966a)
65 were ever
of
were incomplete,
counts
For example,
site.
a group
in
Most
impractical.
made censusing
which
under conditions
Fable's
Anecdote's
troops
troop
contained
in
were smaller
(Kolmogorov-Smirnov
of troop
the
two-sample
3.2
about
about
number of animals
for
constant
contained
troop
the
size
at
least
could
dry season
than
test;
p<0.001
they
4 weeks
be detected.
(median
250 individuals
135 (median
in
count
22).
were in the
for
Fable's
0
3.1:
Figure
frequency
Cumulative
made on two troops
with
in two
which counts
seasons.
100-
increasing
size
0
8
02
O"
N
UO
"
o"
O"
0O
aV
50
ö
.4r4o
of
"
r,
v
O
.
Season
"
4
O
rz 0
Troop
Wet
Fable's
C)
"0
Anecdote's
10-20
50-60
90-100
Animals
of
-Number
Dry
Q
120-130
in
Count
150-180
were
troop,
for
p<0.05
dry
the
both
In
seasons,
but
animals,
Separated
troop,
large
were
(about
fragment
set
from
off
the
Both
troop.
hours
Occasionally
for
rejoin
Fragments
at
behaviour
they
for
morning
remained
1.5km, and were
before
about
450-500
on the
evening
away.
During
censused),
third
this
(Southern
hours
in social
behaviour
three
and did
not
during
time
in
and
(Fable's
troop)
Break's
troop).
(Lucky
troop
them, spent
approached
they
were
(453,496).
of
together,
before
at widely
3.3
other
moving
separated
in
the
They
moved about
social
behaviour,
(425
site
then
two troops
together
2km
individuals
In the morning,
vantage
together
from about
site
in
seen
two troops,
a sleeping
off
where
they
two hours
once
They shared
and the
site
site.
These
least
censused
in social
was once
a sleeping
137 individuals).
south-west,
observers
at
On
known troop.
sleeping
which
resting
always
not
sleeping
troops
twice
moved off
sites;
of 275 formed
known
nearest
feeding,
the
a group
season
2 hours,
two
troop,
the
tkm.
engaged
site,
afternoon.
time
to
point
the
as they
troop
At this
dry
members between
8/8/78
of
over
but
largest
in the early
the
and counted
troop
moved to
from
another
with
two
sleeping
separate
the
than
and then
foraging,
of
coalesce,
a sleeping
the
about
again
The larger
with
4km
seen
fragmenting
association
In
for
together
for
by just
frequently
200m from
30 min,
about
the
from the main troop
larger
groups
overnight.
remained
late
form
roughly
to
observed
of 311 formed
a group
of
day.
also
to
sites,
sleeping
29/9/77
were
at
to
remainder
been separated
a
on 27/7/78
independently
foraged
they
13 became separated
the next
them until
of
spend the night
might
of
a party
135) was observed
having
each other
rejoined
For example,
own.
the
troop.
the
members of the
of other
independently
followed;
were
the parties
example,
(of
main
7
few
as
as
or more of
on their
completely
site
sleeping
might
earshot
within
50) of the troop
fragments
and then
foraged
the
away from
include
a third
made up of
moved off
occasionally
break
These parties
occasionally
normally
parties
but
parties
probably
3.1).
frequently
would
independently.
and forage
troop
(Fig
troop
season
in
troops
both
for
count
dry
typical
50 and 90 individuals
between
contained
17, and the
was about
season
The median
troop).
Anecdote's
a
with
Southern
spent
two
to the west.
points
(Whiten,
Hall,
Sharman)
(n=5).
them (465,412,424).
counted
The composition
given later
in this
this
of
(section
chapter
3.3.1: Number and Population
A lower
limit
collected
single
sleeping
Fable's
troop's
7.2 baboons
6.
in chapter
Fable's
troop
similar
densities,
around
8.7 baboons
shared
about
This
of
size
home range
than
but
were counted
was 55 (n=16;
the
range
(dry
the
within
of
skewness
= 0.007
the
of
genus
are
log-normally
2.60
ns),
=
within
each
species
P. papio
(data
from
significantly
P. hamadryas,
species
than
smaller
but
(Table
is in any other
larger
3.1).
troops
Papio
troop
of
for
size
known for
Dunbar
than
Variance
Papio species,
dry season
is
also
this
the
in
size
log
al.
et
sleeping-group
troop
size
for
P. papio
so for all
of
48.4
mean of
The mean log
Boese
all
is
of
(n = 179;
distributed
and
park
sizes
log-normal.
mean log
3.4
Troop
troops
the
The distribution
mean
sigificantly
P. papio.
troop
larger
prep).
study
in troop
in
elsewhere
a transformed
with
and Sharman
the
The median
park.
The typical
3.7).
sizes
(data
size
at
home range was
in the wet season were therefore
troop
individuals
sizes
the
troop
ns; kurtosis
from
in
elsewhere
troops
season)
range
baboons
baboons
Papio species
10-236) (Table
and the study
median
If
troops.
other
of this
density
5.5 and
between
between
with
baboons
of
on home range
gives
overlap
must
per sq km.
troops
at Mt. Assirik
for
the mean occupation
then
data
from
comm)
Assirik
density
on the
calculation
its
3.3.2: Comparison of P. papio with other
Sixteen
limit
about
(pers
Mt.
a
shared
same night,
at
data
from
and S. Harrison
population
accounting
half
is
census,
individuals
On the
can be calculated
sq km, without
per
The
An upper
home range
area to be presented
Valley,
at Lion
770.
than
630
(Map 6.1).
Valley.
Stella's
be greater
therefore
in
in
single
be obtained
could
when about
Valley
the night
spent
baboons
8/8/78,
of
the
of Baboons at Mt. Assirik
population
at Secret
site
140 individuals
heard
night
total
to
according
3.7.1).
Density
on the
on the
group,
450±33
was
The mean count
of
troop
1975)
is
size
of
other
larger
but P. ursinus.
Papio
than
it
Table 3.1:
Comparison
Mean Ln Troop
of
Size
Papio
of
species
Significance
level of
difference
between
P. papio and other spp.
(t and F tests)
Species
n
mean
sd
mean
P. papio
{ 39
4.23
0.75
P. anubis
P. cynocephalus
P. ursinus
59
1 47
1 25
3.59
3.89
3.68
0.33
0.52
0.58
--
<0.001
<0.025
<0.005
9
8
4.59
4.06
0.34
0.21
<0.05
ns
P. hamadrxas
P. hamadryas
sleep-group
bands
densities
Population
(P. anubis,
n=
The
density
population
baboons for
3.4:
of
the
the
same social
(Chapter
A troop
moves in
the
troop
members for
This
dry
segments"
definition
of
the
been
not
they
tenuous
at
the
that
more
a cohesive
certain
social
for
unit
a relatively
wet
at
used
because
Break's
troops
were
known
that
individuals
bonds
to
part
of
it
is
troop
another.
Members
day.
from
part
members
one
in
the
not
were
in
distance
small
social
exchange
individuals
with
least
share
other
troop
day.
means that
called
of
to mean a group
apparently
which
in
forage
are
and
network,
best
a
at
population
all
use "troop"
I shall
and Lucky
is
living
Discussion
has
network"
it
10.3 km-2;
density
was probably
Fable's
most of
seasons
together,
seen
2
km
to 25.1
Kummer 1968)
example,
while
have had at
would
of
9),
thesis
this
"social
for
whether,
of
are
The term
clear
of
which
animals
bonds.
remainder
highly
are
is known.
at Mt. Assirik:
Group
Papio
than the median for
lower
which the figure
The Social
In
at
slightly
Assirik
Mt.
<0.001
<0.025
ns
<0.025
<0.001
genus
(median
1971)
et al.
population
the
of
(P. hamadryas,
1.8 km-2
Aldrich-Blake
13).
baboons
of
from
ranging
variable,
1
sd
the
"troops".
season
"group"
3.5
social
Dry
groups
season
(Carpenter
seen
troops
in
both
wet
may be
and Nishimura
and
"group
1969),
or
may be "clans"
they
is
there
from
in
no advantage
identity
the
of
organisation
the
making
either
assumption
baboons at Mt. Assirik
is
1976),
the
until
better
group is discussed
of the social
and nature
(Abegglen
season "bands"
wet
but
social
The
understood.
further
in Chapters
10 and 11.
Group
but
species
groups
groups
of
Papio
of
P. papio
P. hamadryas.
baboons
to
tend
The cause
of
is
partly
be larger
than
such
a
species
those
of
any
is
not
differences
species
known.
Baboons
troops
cohesive
and
baboon
were
in
exceeded
bands
baboons.
of
comparison).
The
that
by
are
much
(Data
from
herd
study
was 370.
about
350 individuals
smaller,
Dunbar
P. papio,
four
herds
and
of
and the
in
known
social
(Dunbar pens comm).
group
autonomous social
since
units,
they
in
the
(Dunbar
they
did
sense
and Dunbar
fragment
season (Iwamoto 1975).
3.6
troop
of
any
whose
size
is
being
(although
bands
to
relevant
this
prep).
519
size
individuals
in
recorded
of which the largest
Although
that
their
was of
the band is
not
a
P. anubis
troops
and
being
assemblages
of
1975),
for
non-
any
P. cynocephalus,
of
T. gelada
may be more coherent
not
in
baboons,
of
contained
herd
median
formed
Break's)
known
more
are
and Sharman
T. gelada
of
recorded
least
at
season,
recorded
troops
not
probably
Mt. Assirik
at
Herds are made up of bands,
P. hamadryas bands are
Mt. Assirik,
of
and four
1975),
and Dunbar
all
troops
ten
troops
wet
largest
other
any
is
and Lucky
among the
two
90% of
the
(Fable's
than
The
in
to
related
probably
Assirik
the
within
well
are
are
Mt.
at
However,
were
larger
than
largest
coherent
at
which
densities
habitat
Mt. Assirik
P. hamadryas,
T. gelada
(Dunbar
for
species.
size
Since
the
were
larger
densities
at
1975),
groups
primate
known
three
of
live
sites.
baboons
social
savanna
other
favourable
unusually
human
at
(Hladik
resources
two
Mt. Assirik
at
those
of
range
of
in
size
characteristic;
yet
Discussion
Group Size:
3.4.1:
long
than
periods
the
troops
in
the
at
dry
At
the
size
group
eastern
mean
encountered
P. hamadryas]...
1968a).
observed
party"
at
at
no doubt
that
P. hamadryas
Mt. Assirik
larger
than
in
interactions
Large
(1970)
Altmann
largest
recorded
of
assemblages
these
troop
discussed
are
Seasonal
paniscus
1966),
for
1980a, b).
Troops
fragmented
into
Kummer
reassembly
the
population
study
site
of
at
to
north
social
one-male
group.
Altmann
and
remains
the
Casual
larger;
far
are
Mt.
This
at
Assirik
baboons
and
(350
south
the
Mt. Assirik.
were
extending
to
of
the
3.7
it
does
Firstly,
the
part
of
from
River
Gambia,
field
the
(pers
obs of
site).
new
adopt
in
a
in
the
near-continuous
to
the
on the
baboons
Even
to
generalise
baboons
a
the
for
account
probably
north
the
nor
to
by
has
resources
scattered
fails
season,
to
rapidly
explanation
these
of
isolation
reproductive
of
Africa,
P. hamadryas
for
individuals)
wet
Some
season.
described
(Anderson
South
in
have
1975),
baboons
Papio
Ateles
1976),
(Hladik
senex
eg
primates,
other
(Homewood
dry
exploitation
in
known in
that
speculates
the
baboons
Niokolo-Koba
the
that
most
recorded.
which
of
the
groups
one-male
groups
to
to
seems
units
which
Suikerbosrand,
at
in
population
of
River
seen
population
groups
the
distribution
the
been
galeritus
P. ursinus
environment.
of
the
yet
a
of
There
social
animals,
and Presbytis
one other
Anderson
adapted
predator-free
200
in
aggregation
baboons.
savanna
though
size,
1980),
smaller
small
behaviours
the
of
(1968a).
this
(Kummer
T. gelada,
baboons
Cercocebus
only
resembled
allowed
group
Roosmalen
been described
subgroups
in
changes
(van
party
below.
(Crook
T. gelada
in
[of
P. hamadryas
of
within
parties
was 275
Mt. Assirik
cohesive
place
have
baboons
savanna
individuals"
Like
take
savanna
of
494
sleeping
in
live
almost
of
"moving
630 individuals.
about
known
one
largest
the
of
areas
aggregations
P. papio.
bands
other
P. hamadryas
largest
mean
where
in
at
sleeping
the
of
also
The
P. papio
for
survey,
was in
comprised
sometimes
those
are
known
227.
was of
P. hamadryas
troops
was
1968a);
seen
far
individuals
The
midday.
P. papio
are
of
figure
broad
it
than
midday
at
(Kummer
750
reach
may
day
the
The equivalent
"moving
larger
number
during
(1968a)
Kummer's
of
was considerably
the
survey,
end
if
the
south,
roads
study
was a part
probably
is
later
discussed
in
this
The observation
that
does
help
rejoined
not
foraging
parties
(DeVore
troop
baboon
have
seen
Aldrich-Blake
Altmann
1970
1968a,
Stolba
T. gelada,
P. ursinus,
1979;
P. papio,
al.
of
1971,
for
Altmann
the
or
rare
sleeping
for
sites
of
Papio
Washburn and DeVore 1961, Altmann
and Saayman 1970; P. hamadryas,
at
Mt. Assirik
seems more like
of
hybrid
T. gelada
or some species
Nishimura
1969, Macaca silvans,
certainly
visibility
(Aldrich-Blake
contact
in
Gautier-Hion
press,
and feeding
(Anderson
1980b,
1978, Williams
other
than
1964,1966a;
1970;
Papio
Hamilton
and
1974).
it
is
at waterholes
P. cynocephalus,
Stoltz
P. ursinus,
In this
respect
baboons
Deag and Crook 1971).
competition
1975),
Dunbar
and
of macaques (eg Macaca fuscata,
3.8
such temporary
explain
Kummer 1968a).
other
Kummer
troops
and Altmann
the
and
have
to associate
Rowell
Altmann
study;
Byrne
(Dunbar
(P. anubis,
a day
of
this
Poor
1970,
troops
space
P. hamadryas,
to
groups.
T. gelada
of
species
1979).
forward
of. subgroups
of
1972,
Pi 1968, Kummer 1968a),
formation
herds
apparently
(Sugawara
body
Nathan
as
1963a, King 1975, Rubenstein
Watt 1970, Hall
Unlike
and
of
reports
every
the
1980b;
1975),
and Altmann
of
in
group members to keep in
1970, Jones and Sabater
may promote
troops
social
main
P. cynocephalus,
Anderson
social
the
and
split
various
early
from
off
reform
1971;
Dunbar
the
Since
1962a),
and
al.
baboons
primate
difficult
Hall
have been put
Two suggestions
fragmentation
breaking
Dunbar
x P. anubis
on group size
sometimes
between
by baboons.
fragment
et
and
of predation
at Mt. Assirik
distinguish
1965,
37;
p
Dunbar
P. hamadryas
et
to
to
the
Secondly,
Chapter.
the troops
and Hall
been
(P. anubis,
makes it
the effect
P. cynocephalus
of
1000 animals.
least
at
known to be possessed
organisations
the
contained
was not predator-free;
environment
the deme (Mayr 1963) of which it
isolated,
was reproductively
population
than
P. papio
it
Carpenter
does
and
Age-sex Structure
3.5:
The demography of
individuals
which
lead
to
1979)
in
in
and
any
demographic
be
the
influence
If
(as
population
not the birth
the
the animals
conditions
file
single
all
of
of
the
the
censuses
difficult
to
Altmann and Altmann
animals
forced
the
broader
it
category
was not
were
in
areas
the
Furthermore,
of
whether
in recent
or declining
such
of
Large
of
good
to make rapid
observer
was doubtful
to classify
than older
at all,
I
in
of
groups
bunching
study
of
the individual
1966,
of the
age and sex.
was classified
juvenile
are
animals
(Crook
visibility
that
felt
and most
rarely,
evaluations
the individual
rather
ideal
Under
a way that
occurred
In many cases in this
1970).
(eg juvenile,
possible
should
we
away from the observer
This
seen.
censuses.
even
Whenever an evaluation
in
clearing,
were partial
census
1977)
possible.
whenever
were moving diagonally
present
animals
expanding
rapidly
Method
censused
a grassless
across
1967b).
Rowell
census of the baboons in the park.
of Troops:
were
between
may indicate
increasing,
will
can be examined by consideration
of any trend
in an earlier
troops
The
a
ratio.
infants
of
classes
to
the
of
by members of
1976,
Verschuren
and
female
adult
three
Age-Sex Structure
3.6:
to
has been steady,
rate
data presented
Dupuy
by
asserted
in
belonged
cannot
strategies
mating
and Dunbar
Rowell
ratio
sex
experienced
in
(Dunbar
adult
may
and Leland
Struhsaker
the
study,
competition
Assirik
Mt.
at
The duration
years.
this
1963,
knowledge
a
1966a,
differences
to
immature
numbers
relative
to
same species
is
high
a
expect
interest
(Rowell
of within-sex
troops
the
troop
may lead
the
of
populations
the
in turn
behaviour
Social
without
in
environment
1979) and this
1979).
interpreted
of
social
(Kummer and Kurt
(Dunbar
correctly
amount
and
group,
and Altmann
behaviour
structure
the
organisation
social
Of particular
1979).
the
(Altmann
in
social
case
influences
a troop
live
variations
Introduction
of Troops:
male),
into
a
and if
was counted
and
scored as an "unknown".
The age-sex
composition
of more than 50 animals.
of
troops
The censuses
3.9
were
estimated
were weighted
from
all
censuses
by summing within
age-sex classes
It
or
across
was always
categories
(black,
To
the
assess
dorsal)
estimate
but
coat
not
could
(1980)
has
the
calculate
Assirik
and Amboseli
to
the
compare
be
fixed
a
at
baboon
troops
the
in
baboons
9/4/78,4
trip
Baldwin
the
roof
seen
the
the
troop
watched
binoculars,
driver,
vehicle
troop,
at
them
to
possible
three
age-sex
Mt.
mortality
at
it
is
possible
of
a stable
that
with
took
part
in
other
for
baboons.
was usually
onto
age-sex
a
To
3.10
the
structure,
(the
at
a good
take
the
in
The
3/4/78
time
one
vehicle,
a
sat
were
point,
and
the
observer
through
a progression,
of
driver
on
baboons
vantage
the
and
For
observer)
census
of
survey
survey
Every
categories
checksheet.
the
(1975).
et al.
person
moving
that
between
out
drove
the
to
a road-strip
was carried
by Boese
unlikely
or
size
censused.
possible
is
possible
also
twice.
the
was
abnormal
of
One person
if
it
demographic
the
that
were
survey
while
least
out
the
any one time
at
groups
us.
which
in
out
point
site
park
vegetation
and called
who noted
the
was stopped,
censused
the
in
15kph,
the
make it
which
Since
a similar
about
scanning
(1979)
The survey
accompanied
at
position
whose riding
Mt. Assirik
at
and myself,
after
years
Land-Rover,
in
Park.
the
used
were
1980 p 19).
lived
Hall
Two people,
or
female)
adult
among infants,
characteristic.
elsewhere
(ventral
an
age-specific
similar
Rowell
Mt. Assirik
baboons
categories
The results
infants
that
any one field
species
I
Troops
Other
at
(p 34)
of
rate
(Altmann
and
observed
structure
birth
with
age
always
an infant
of
be placed.
should
infant
coat.
data
was roughly
(1970a)
Crook
(nearly
By assuming
recent
Comparison
3.6.1:
of
the
proportions
baboons
of
population
three
infant's
provided
Amboseli.
at
the
three
the
of
be recorded.
relative
classes
in
age category
probable
colour
Altmann
infant
it
the
of
colour
the
position
carrying
animal
the
the
which
brown)
infants
of
possible
the
with
or
dorsally
was riding
into
decide
to
always
proportions
an infant
say whether
transitional,
the
on
together
to
not
whenever
recorded
to
possible
but
ventrally,
censuses.
animals
was
to
the
sometimes
to
able
troop
the
watch
observer.
Whenever
vantage
for
point
(since
road
possible
taken
each,
of
each series
baboons
the
and to
years,
The
from
the
time
each age-sex
each
class
age-sex
Table
different
in
3.2.
In
in
one troop
of
was the
no case
from
4
previous
for
described
are
with
with
three
declining
was
which
many
compared
especially
and
to
females
adult
of
Mt. Assirik
its
in
included
class
14 largest
the
age-sex
trends.
been
from
the
the
over
park
have
The
troops.
within
in
two
and
Results
The proportion
censuses
changes
the
of
censuses.
of Troops:
across
the
with
compared
census
and
one
Age-Sex Structure
of
in
baboons,
of
the
of
census
earlier
immatures,
data
The
examine
ratio
sex
P. cynocephalus,
of
at
adults
an
population
of
to
studies
in
baboons
of
adult
other
increasing
for
censuses
proportion
for
is
given
any age-sex
class
the
troops
both
mean proportion
in
deviation
standard
mean and
of
varied
census
any
other
(t
troop
test,
p
> 0.05).
always
The
between
p<0.025
only
two
the
variability
from
troops
and black
in
in
differences
transitional
data
the
quality
quality
also
were
study
to
was used
baboons.
of
populations
3.7:
of
field
(1975)
indications
provide
highest
as
analysis.
al.
This
the
was made of
the
of
the
leave
not
As many censuses
park).
and a record
this
et
population
and
populations
those
Boese
proportion
immatures,
the
during
park.
the
of
structure
in
by
collected
in
in
collected
the
The results
scale.
were used
The data
in
a better
to
moved
we did
although
censuses,
permitted
then
the
by
classifications
was
vehicle
on each group,
on a 5-point
of
those
possible
was not
check
the
subsequent
this
were
a
as
the
on
every
in
both
were
infants,
case),
which
in
difficulty
troops
the
for
the
variances
among
and
were
seeing
analyses.
3.11
younger
infants
the
(F
ventrally
subject
As a result,
classes
independent
juveniles,
carried
classes
them.
age-sex
within
to
the
I
test;
greatest
pooled
the
Table 3.2:
Proportion
in the
various
age-sex classes
censuses for two troops
of
(n=14)
Fable
Class
Male
Mean(t)
24.26
sd(%)
sd(%)
est
3.21
61±8
22.30
3.95
30±5
2.42
1.59
2.16
1.52
81±14
28.41
6.33
43±9
1.91
7±5
3.50
2.18
7±3
4.47
1.73
27±11
37±4
11.17
14.70
5.47
4.49
15±7
20±6
1.43
1.12
Female
29.20
5.70
Subad M
2.97
Older J
Young J
10.72
14.24
6.02
3.78
Trans I
Black I
0.37
0.27
0.69
0.56
Ventr. I
Dors. I
2.76
4.78
1.37
2.23
note that
est
6.11
4.83
0.86
0.42
1.24
1.13
3.79
3.84
2.93
2.65
100
age-sex
class
numbers
in
Structure
of
pooled
50 animals
(small
(large
censuses).
animals
in
these
deviation
censuses
of
in large
Wilcoxon
therefore
animals
censuses),
in
in
Table
35 (45%) included
(55%)
3.3,
censuses was smaller
more internal
were recorded
general
matched
signed-rank
of
fewer
50 or
included
the
with
more
class
each age-sex
together
than there
influenced
of
the
by
than in small
pairs,
standard
censuses
There
p<0.01).
in censuses
self-consistency
The mean proportion
not
and 42
The mean percentages
given
troop
each pe-rcentage.
Variance
tailed
are
in
Population
the
Out of 77 censuses of more than 20 animals,
than
20±7
2.
sum of mean % is not necessarily
= estimated
Age-Sex
3.7.1:
36±9
code see Chapter
age-sex
(n=14)
Anecdote
Mean(%)
2.04
1.02
KEY: for
complete
est
Oestrus
Pregnant
Brown I
14 most
(twowas
in which 50 or more
was in censuses of fewer animals.
census
the
3.12
made up by each age-sex
number
of
animals
in
the
class
was
census.
Table 3.3:
More or Less
Containing
Censuses
{
small
(>20 but <50)1
n=35
%
sd
23.0
8.6
1 21.7
4.6
Oestrus
Pregnant
1.6
1.3
2.6
1.9
1 2.3
1.0
2.1
Female
30.4
9.5
1 29.3
1.3
5.9
Subad M
3.9
3.3
3.1
2.2
Subad F
0.3
1.2
1 0.1
0.4
10.8
6.6
1 11.7
5.0
Young J
15.3
6.2
1 16.9
6.6
Brown I
Trans I
Black I
Ventral
Dorsal
2.9
0.1
0.1
3.0
6.2
5.2
0.5
0.6
2.7
3.8
1
1
1
1
1
4.8
0.9
0.7
2.2
2.4
Male
j
Older
KEY: for
was
in
seen
one
small
following
large
(>=50)
n=42
sd
%
Class
There
50 Animals
than
I
than
analysis
have
large
in
pooled
ones
data
infants
transitional
independent
fewer
censuses
2.
code see Chapter
age-sex
exception:
5.1
0.5
0.2
2.9
4.2
(t
test,
from
For
the
animals
or
p<0.025).
50
of
censuses
were
more.
The age-sex
by
the
fragmentary,
as
shown
proportions
larger
censuses
A
censuses.
of
between
them 4977 animals.
1: 1.42.
and infants
data
are
states,
Subadults
14%.
shown in
better
estimate
Of these,
and 23% were
adult
made up 3% of
The adult
detail
female
in
is
Table
3.13
the
to
3.5,
of
assessed
not accurately
since
100 animals
than
The 42 censuses
results
reproductive
censuses.
3.3,
Table
in
less
of
is
of the population
structure
are
the
given
by
provided
more
32% were
males.
censuses,
immature
and are
same weight
pooling
adult
The adult
juvenile
ratio
taken
in
females
sex
various
ratio
animals
was 1: 1.31.
to
the
included
50 animals
than
more
perhaps
smaller,
represent
was
28%,
These
the
approximate
the
during
Assirik
the
of
proportions
Summary of
Censuses
Specific
Age-Sex
Class
Total
Male
1130
22.7
Oestrus
113
2.3
1
3
Pregnant
Ventral
58
157
1.2
3.2
0
2
2
4
Dorsal
234
4.7
4
7
Female
1037
20.8
Subad M
151
3.0
2
4
Subad F
5
0.1
0
1
Older J
Young J
582
11.7
10 14
759
15.3
13 17
37
0.7
0
287
23
13
157
5.8
0.5
0.3
3.2
58
02
02
24
234
4.7
ü7
Brown I
Trans I
Black I
I
Ventr.
I
Dors.
Summary
Age-Sex
Class
binomial
%
95% cl
of
20 26
19 24
2
group
juveniles
and
censuses
on the
From data
Fable's
group
76
Mt.
Male
1130
22.7
20 2
Female
1599
32.1
29 35
Subadult
156
3.2
Juvenile
1378
27.7
25 31
714
14.3
12 16
infants)
study
on page
133
similar
groups
in
this
Chapter,
groups,
can be calculated
(Table
The
3.3).
squared
and Anecdote's
of
the
= 7.40;
df
= 4;
the
3.14
probable
troops,
of
males,
subadult
that
to
two
composition
18
and a hypothetical
3.5).
made up of
group,
(chi
presented
24
separately.
females,
adult
was
binomial
95% cl
%
large
(reported
males,
adult
at
Total
Infant
an exceptionally
was made on 29/9/77
(98
troops
50 or more Animals
of
KEY: for age-sex code see Chapter 2.
In this table, V. D and F are listed
A census
both
study.
Table 3.4:
Juvenile
in
classes
age-sex
population
this
100
of
ns).
composition
'standard'
dry-season
of
Table 3.5:
Possible
Composition
of Troops at Mt. Assirik.
Adult
Subadult
Males
Females
Fable
Anecdote
Dry
57
31
80
43
Season
20
29
Troop
Juveniles
Infants
8
4
69
37
36
19
3
25
13
Males
Infants
3.7.2:
The age and position
infants
Of
recorded.
the
these,
(46%) and black
dorsally
riding
(Table
(27%)
ventrally
riding
were
infants
were brown
classes
principal
313 infants
of
3.6).
Table 3.6:
on Adults
Riding
Infants
Age Class
Position
on Adult
I
I
I No.
i
184
Ventral
%
26.8
142
i
Dorsal
ITransitional
Black
13.4
1126
transitional
to
the
survived
categories
category
were black,
for
of
percentage
172.2
I
I
1 30
1
should
4.5
two
led
to
1
98
1
1
215
1
1150
then
that
might
a
expect
(Altmann
successively
3.15
the
et al.
lower
infants
are
P. cynocephalus
spend
27.0% of the
If
percentages
to
313
that
and 63.5% brown.
8.3% and 58.3%)
have
Total
=
3.4 and 3.6 suggest
months,
life
its
%
1143 45.7
1
1 37
infancy
throughout
No
i72.2
9.4% transitional,
about
(33.3%,
%
No.
The combined data from Tables
infants
Brown
are
very
close
infant
in
1977).
percentages
each
Deaths
being
which
the
of
in
any
found
in
the
three
tallied
categories,
closely
in
spend
were
However,
Boese
59.9%
the
of
but
the
3.7.3:
the
the
Censuses Elsewhere
(Table
troops
3.7).
season,
they
when,
represent
censuses
more
Of
an
infants.
brown
8.3%,
32.2%,
and
transitional
the
by a similar
either
that
recruitment
rate
1972,
since
and
40%, 8%,
about
population,
At
both.
or
at
ratio
of
were collected
troops
tended
males
them
and
Subadults
infants
9%.
be taken
The 9
of
records
34% were
the
be smaller
to
GPN de NK.
the
between
in
therefore
should
in
P. papio
1: 1.54.
and
survey
data
included
adult
of
31%,
least,
These
survey
road-strip
road-strip
sizes
22% were
the
Park
the
of
990
females,
adult
made up
4% of
the
adult
female
to
The
was 1: 1.18.
proportions
the
of
similar
in
these
(Table
3.8)
with
proportion
(Cochran's
the
50 animals
juveniles
ratio
larger
in
wet season.
sex
the
suggest
that
or
declining
censused
from
than
in
categories.
troop
these,
adult
population,
immature
the
minimum
of
animals.
giving
in
were
they
has been falling
Mt. Assirik
at
infants
in the Park
were
The data
is
may be affected
1974,
in
years.
It
of
black,
were
unbiassed,
slightly
three
similar
data;
data
of
low
park
or
numbers
observed
are
was also
a stable
into
Sixteen
The
figures
in
population
and 52% fell
they
The two sets
infants
in
provide
Mt.
black
the
to
mortality
at
two
of
numbers
relative
infants
the
of
Amboseli,
to
al.
289
if
mortality
than
et
the
that
(1975)
respectively.
bias,
dry
likely,
to
likely
were
infant
last
the
over
rate
underestimated
censuses
of
falling
birth
been low
may have
each category
infants
time
of
in
observed
numbers
length
relative
category,
perhaps
possible,
the
since
there
a
or
brown
the
with
each
Assirik,
but
censuses
to
censuses
the
of
the
exception
the
adaptation
made
infants,
of
the
at
each
which
Mt.
appeared
to
than
t
test
were
Mt. Assirik
from
Assirik
Fisher-Bihren's
class
age-sex
figures
corresponding
population
of
by
up
make up a
elsewhere
t'
_ -3.26,
p<0.01).
Boese et al.
adult
males,
(1975)
30% adult
1723 individuals,
counted
females,
9% subadults,
3.16
which . 16% were
29% juveniles,
and 17%
of
Table 3.7:
During
Road-Strip
Censuses of Troops of Baboons Encountered
du Niokolo-Koba
Survey of the Grand Pare National
Best
Age-Sex Category
S0Y
I
U
n
M
F
1
2
3
24.2
20.2
22.8
32.2
37.6
36.1
4.2
2.2
5.1
13.6
18.0
9.5
19.5
12.4
15.8
6.4
9.6
7.6
0.0
0.0
3.2
236
178
158
4
19.2
36.7
5.8
12.5
10.8
15.0
0.0
120
5
6
16.7
29.0
27.8
30.6
3.3
1.6
23.3
6.5
20.0
6.5
8.9
8.1
0.0
17.7
90
62
7
16.1
32.1
1.8
17.9
21.4
10.7
0.0
56
8
32.1
17.9
5.4
17.9
19.6
0.0
0.0
56
9
13.0
38.9
1.9
14.8
20.4
11.1
0.0
54
10
11
12
34.0
15.0
21.1
8.5
37.5
23.7
4.3
7.5
7.9
8.5
17.5
26.3
10.6
10.0
18.4
2.1
12.5
2.6
0.0
0.0
0.0
47
40
38
13
23.5
26.5
5.9
26.5
17.6
0.0
0.0
34
14
17.2
20.7
3.4
17.2
37.9
3.4
0.0
29
15
46.7
13.3
0.0
13.3
13.3
6.7
6.7
15
16
10.0
40.0
0.0
40.0
0.0
10.0
0.0
10
Census Number
Table 3.8:
Proportion
of various
(estimated
from censuses
Age-Sex
Class
%
Boese et al.
Road-Strip
At Mt. Assirik
%
95% cl
95% cl
%
95% cl
M
22.7
20 26
22.1
19 25
15.7
14 18
F
32.1
29 35
34.0
31 37
29.8
27 33
s
3.1
2 4
3.8
3 5
8.5
7 11
0
Y
11.7
15.3
10 14
13 17
14.8
16.4
13 17
14 18
12.4
16.9
10 14
15 19
I
14.3
12 16
8.8
7 11
16.8
15 19
n
4977
990
age-sex
code see Chapter
KEY: for
infants.
age-sex
containing
in population
classes
50 or more individuals)
Their
data
are
compared
3.17
with
1723
2.
the
data
from
this
study
in
3.8.
Table
They
significantly
infants
more
troops
the
of
adult
adult
animals
adult
males
Table
were
They
males.
in
at Mt. Assirik
the
at
genus
number
males
than
of
per
did
adult
fewer
significantly
recorded
troops
per
(Table
female
than
of
troop,
number
subadult
the
Although
the
more
of infants
proportion
other
found
the
in
immature
most
the
other
female,
road-strip
lay
and
of
adult
female,
adult
range
in
and
the
proportion
baboons,
with
immature
animals
per
3.9).
3.9:
of
Demographic
parameters
to the
baboons relative
of known populations
at Mt. Assirik.
population
Demographic
Parameter
I
for
mean for
mean
Iall baboonslgenus Papio
Proportion
I
I
S 0.46±0.08
adult
of
animals
Number of immature 1 1.69+0.58
animals
per ad. fem. =
Number of adult
males per ad. female{
Mount I
Assirik{
I
I
{ 0.47±0.08
II
I
I
II
x(n)
p
0.54
{ 28(33)
0.16
i1 1.66±0.57
1 1,31
i1 26(35)
0.23
I
II
1 25(35)
0.17
I1 0.45±0.23 I10.47±0.24
II
I
i
ii
i
0.70
KEY:
x
number of known populations
of baboons in genus Papio with a
less extreme value than that found at Mt. Assirik.
(n) number of populations
of baboons in genus Paplo for which the
value was known
in genus Papio with a more extreme
p
proportion
of populations
value than that seen at Mt. Assirik.
3.18
of
animals
of
populations
the
but
figure.
the
had a higher
they
fewer
their
within
proportion
per
significantly
survey,
with
well
animals
female,
adult
reported
corresponded
Mt. Assirik
Papio
also
and
males
adult
of
more
adult
3.8:
Age-Sex Structure
of Troops:
The
the
in
lower
Fable's
The only
were
these
categories
classes
P. anubis
infants
even
obs),
the
while
Comm).
riding
rather
more timid
infant
numbers
alarmed
female
the
Mikumi,
or
the
troops
the
were
of
than
result
of
(pers
(Lee
pers
infants
were
troop
seemed
in
variance
and easily
visibility
poor
been
declines
high
the
a
and
alarmed
Anecdote's
so that
have
humans
likely
troop,
so that
ventrally
presence
a census.
Fable's
that
transitional,
riding
more
were
in
of
baboons.
for
each
Ruaha
The population
(Western
and van Praet
there
equivalent
this
figure
if
in
the
baboons
of
juveniles
comparison
with
other
(1980)
is still
in
of
mortality
1.9
and Altmann
It
is
the rapid
has indeed
to
and Oliver
each adult
1970).
The
therefore,
growth referred
to by
to decline.
is
females
been low
rapidly
unlikely,
may have started
at
immature
was declining
Amboseli
(Altmann
known
rapidly;
(Lee
was 2.3
figure
immatures
3.19
were
per
comm) felt
1980) at a time when for
animals
and it
(pers
expanding
there
population
undergoing
(1977),
and Lee
were
studying
comparable
1973, Altmann
proportion
infant
the
few
in the GPN de NK was 1.3.
population
low
were
were 1.4 immature
Dupuy and Verschuren
The
they
female
adult
at
Rasmussen
Both
that
populations
by
animal
rather
with
a population
adult
1979).
interpret
infants
to
missed
did
the
when
states
should
black,
each of
would
6 weeks,
about
of
numbers
Owens (1972)
Mt. Assirik
at
in
in
fluctuations
However,
ventrally
per
while
that
of
indicate
animals,
female
infants
three
infants
of
numbers
at
two troops
categories,
age-sex
dorsally
baboon
have been
might
four
Furthermore,
habituated
be
than
censuses
populations
that
the
troop.
animals.
The
adult
more
to
of
ventrally
riding
dorsally
censuses
and small
the
proportion
Infants
the
observed.
were
rode
become
in
ride
of
than
infants
troops
to
proportion
brown
Fable's
variance.
started
dorsally
of
in
a
presumably
censuses
Absolute
low.
troop,
was
class
age-sex
in
variance
very
every
included
noted
infant.
of
large
much greater
riding
the
were
introduced
have
as
differences
was in
which
numbers
of
Anecdote's
in
than
larger
significant
Mt. Assirik
mean percentage
troop
the
of
consequence
at
in
variance
Discussion
in
the
to
difficult
park
for
at
4 years.
least
the
the
study
by
collected
recruitment
it
was during
than
started
Boese
(1975)
al.
et
so that
continued,
four
the depressed
is correct,
interpretation
apparently
that
was higher
the year
can be seen in the data on troops
pattern
similar
indicated
the
proportion
of
in the park
If
in
this
troops
has
them is
now
in these
infants
of
A
the study.
previously.
recruitment
a year
about
elsewhere
years
in
of infants
the age structure
that
seems possible
troops
study
before
It
low.
The age-sex
to
similar
immature
structure
that
Mt. Assirik,,
at
in
represented
of the population
this
indicated
animals
but
study,
many of
this
et
al.
may be starting
classified
these
comm, Altmann
3.9:
Predation
size
There
are
are
poor
size
troop,
and
individual
is
improved
size.
would
hence
to
may benefit
the
baboons
notoriously
(Strum
well
as
by ecology,
determined
at Mt. Assirik
favour
should
Cook and Martin
of being
probability
be
detected
areas
lead
to
difficulty
its
fragmentation.
to
the
argue
that
then large
small
groups,
predicted
that
was good,
since
(1970)
coordination
of
the
With
an
good visibility
Once
Ldetection
of predators
a group,
as the number of prey in the group gets
3.20
(1974)
are
where visibility
in
visibility,
detected
than
and Altmann
in
by belonging
between
relationship
Altmann
would be largest
is
males
group in the wet season.
Firstly,
to
likely
of visibility.
visibility
on
views
has a constant
more
irrespective
group
two
and group
each animal
groups
in a large
living
the individual
of
in
as adult
classified
partly
in the environment
or features
their
in
males
comm).
is
a species
within
ones
with
Introduction
and Size of Group:
group
some feature
if
comm. Popp pers
pers
to
females
adult
adult
and adult
species
other
well
on a major crash.
been
have
subadult
in
observers
pers
predation
of
less
were
whose manes were developing;
animals
probably
Classification
between
If
would
animals
study.
variable
as "subadult"
the park was
by comparison
and fewer
more subadult
recorded
of
ratio
which,
a population
known to have been declining,
Boese
The
sample.
wider
infants
that
except
in
elsewhere
larger
(Bradbury
and
Verencamp 1976b, Cushing
1972, Powell
The
1974, Pulliam
opposite
size
favoured
under
by
prey
(Brock
the
the
of
the
being
1976b,
of
is
in
which
1971,
season,
baboons
as groups
groups
of
changes
made by the
radius
the
I
kept
"wahooing".
Small
troops
generally
too
difference
in following
aware no theory
can
detect
detectability
group
size
to
have varied
in
their
prey
and group
and noise
the
the
group
larger
and the
has
it
chance
of
and Verencamp
If
in
the
not
200m or
little
by
size
as well
will
environment.
3.21
the
group
size,
1974),
Jarman
1976,
but
there
where
the
was
depend on the
accoustic
no
As far
case of
the
follow,
to
difficult
would
males
more when adult
for
by the
detection
conditions,
normal
but
as on the
hand
other
of 100 and 200 animals.
noise,
in
The effective
noise,
been developed
largest
baboons largely
the
extremely
were
the
on
(Estes
vision.
50m under
about
being
of
wet season.
with
contact
by
do
difficulty
by changing
risk
to
in
changes
seasonally,
predation
known
in
a troop
has yet
be
of
of
(Bradbury
to
responded
was about
increase
less
the
was good.
animals,
troop
1972),
Lazarus
itself
size
upon
should
detection
the
of
any one attack
mammals are
occasionally
was
1974,
depends
groups
because
independent
who showed
predator
Large
good.
visibility
have been largest
should
(1964),
1972).
should
to
noise
there
is
Mt. Assirik
visibility
other
group
finds
Lazarus
size
At Mt. Assirik
of
by the
in
when
responded
the
animal
at
group
of
Jarman
by a predator
coordination,
dry
an
baboons
the
by Olson
probably
1971, Lazarus
1975a).
supported
poor
1960,
1968, Hamilton
is
when visibility
predator
Hamilton
If
the
group
conditions
selected
the
detection
and Riffenburgh
group
1973, Treisman
prediction
that
mathematically
the
and Harden-Jones
since
apparent
as I am
that
predators
relationship
relationship
properties
were
between
between
of
the
Baboons
were
visibility
season,
detect
around
in
For
5m of
within
the
dry
On other
cause of death.
whole
The animals
than
female
to
the
example,
beneath
of
with
came
while
Since
changed,
than
season
wet
a
which
also
the
of
and leopards
(but
the
a dormitory
difficulty
of side 2cm of the right
the bone,
to orbit
the
had been badly
frontal
palate
near the sutures
3.22
of
a
of eating
a
in
was
one animal
adult
contusions
She was
her nostrils.
sitting
An
up.
A triangular
fractured.
had broken away
was split,
between the
1970).
A dying
above the orbit
(maxilla)
since
causes other
She had slight
eyes and had great
section
from
death
beneath
her skull
skull)
a
of the
(Brain
by a predator.
tree.
cache of
no hint
with
have died
observed
a sheer-
of
not surprising
the
not
might
These bones
the
for
by
possibly
are capable
of blood
that
an adult-sized
3m x 0.5m top
suitable
12
although
study,
of
were found,
an attack
revealed
ran from orbit
ease
a leopard.
is
found
were
bones
eye and a trace
of
season.
girdle
flat
possibly
were found
result
open her
rest
dry
had been perforated,
on the
fragile,
autopsy
from the
I
had walked,
in this
pectoral
two skulls
the
For
above her right
unable
rather
including
was found
in
was all
season
probably
greater
the size
of about
a site
whose skulls
not
the
from
occasions
is
predation.
probably
the
troop
were observed
That few remains
baboon,
2)
the
certainly
were found
boulder,
baboon skeleton
the
in
that
troop
wet
many baboons
(Chapter
The scapula
a predator
2m high
leopard.
the
the
comm) in
from
of predation
and some baboon fur
sided,
in
been expected
was evidence
until
wet
season.
baboon were discovered.
of
(pers
itself
bones and bone fragments
canine
which
was almost
No incidents
the
near
season
hidden
risk
occasion
have
there
undetected
one
Byrne
to
with
have
and predation
was in
leopard
happened
might
However,
the
with
comparison
normally
might
remain
on
example,
changed
predator
good,
could
by
which,
good time.
a hidden
same thing
visibility
in
(pers
baboons
the
in
dressed
observer
detect
could
observer
season,
a predator
it.
the
a hidden
detect
to
was generally
predators
season
either
it
dry
the
Results
able
before
well
in
and
obs),
the
frequently
clothing
cryptic
to
and Size of Group:
Predation
3.10:
frontal.
and a crack
nasal
and
Her death
maxilla.
was almost
the
certainly
result
from a
a fall
of
beneath.
branch about 10m above the ground onto her head on the boulders
Baboons
reacted
the
baboons
of
noise
two
trees
but
area,
Until
(Rene
the
Bonang,
Predation
The
were
these
of
reactions
dogs
present
in
of
them for
food
baboons
to
the
the
in
baboons
by
seen.
was not
predators
predator
close
wild
of
also
on
observed
from
presence
hunting
Alien,
and were
dogs
with
has
humans
and Size of Group: Discussion
(1965)
interpretation
baboon
of
(Anderson
overstated
on the
influence
clearly
played
1972,
some part
Altmann
Serengeti
in
in
kills
1972).
Harding
1970;
P. ursinus,
Hall
a tiny
of animals
part
of
in
he recorded
which
National
100 food
Leopards
baboons in their
3.23
diets
of
diet
1976;
1963a,
preying
on
large
of
anywhere in
items.
1000 food
over
items.
park was of
the
by lions
saw no baboons eaten
6 baboons in
in Kruger
Schaller
known to include
(1972)
a survey
1956a, b,
few records
rather
form
generally
At Manyara he found
2000 lion
and
(T. gelada,
baboons
Booth
P. anubis,
Altmann
of
has
it
and Pegram 1969, Dunbar and
1967a, Bert
et al.
are admittedly
Schaller
predators.
habits
possibly
For example,
baboons.
of
is
have had a major
may indeed
predation
the roosting
study;
Baboons
baboons.
in
Bert
this
There
but
in the
factor
single
behaviour"
social
and
ecology
organisation
social
P. cynocephalus,
1965b).
remark:
1980b),
Crook 1966; P. papio,
Nathan
between baboons and the
"relations
can prey on them is the most important
that
animals
quoted
the
to
of
precipitately
hyaena
baboons
the
comet.
DeVore and Hall's
the
to
Spotted
people,
pers
flee
man was a major
ago
local
to
reacted
the
of
2km away),
over
(the
leopards
been described.
already
3.11:
They
reaction
15 years
with
area,
pers
and
calls
night-time
obs)
and "wahooing".
the
about
the
in
to a predator.
no resistance
be heard
could
lions.
with
encounters
to
noisily
(Hall
occasions
climbing
have offered
and could
she seemed
when she was found
She was helpless
to have been in good health.
mid-morning,
from her injuries
is unknown, and apart
The cause of the fall
Fewer than
one in
(Pienaar
baboon
a
the
Serengeti
have also
(1 of
164 kills,
Schaller
1969,
been
1972,
2 of
55 kills,
killed
by
Kruuk
leopards.
predation
on
recorded
3 kills
an
and
(1980)
and Altmann
baboon
mentions
(1971)
reported
years,
and Rhine
an
important
(Dittus
1975,
of
wild
the
threat
at
in
probably
and hyaena
seasonal
the
man
in
changes
wet
given
for
worse,
in
visibility,
probably
through
the
had
predation
up a tree
escape
thus
possible,
and
predation
principally
from
ambush
to
a higher
from
the
reducing
man and dog
of
combination
Most
leopards
especially
Since
have led
cursorial
of
predator.
would
quite
certainly
chimpanzees
in
the
Some support
those
pressure
predators,
predation
large
at
differing
1973,
in
and 38.0 respectively)
3.24
this
hypothesis
of equally
the
risks
predation
risk
1973).
while
troops,
Baboons did
so that
at these sites
is
bad,
(Mikumi,
were said
those at Budongo
(Ransom 1971),
Park,
part
Ransom 1971).
Gombe Stream,
in smaller
living
Gombe Stream
predators
may in
under conditions
(Paterson
less
season
for
(Rasmussen 1980),
heavy predation
suffered
of
Paterson
especially
as predators
were few other
areas
and dry
wet
living
troops
Budongo,
Baboons at Mikumi,
(80.2,63.5,
came
changes in group size.
Rasmussen 1980,
almost
man has
of
the
while
avoid,
predation
by data from other
to suffer
in
one troop
2
in
season.
Differential
account
Ransom
ambush predators.
Lions
species.
visibility
(1975)
son".
two species
been
have
a formidable
probably
a
to
so that
normally
easy to
in
primates
and two
was wooded,
would
eating
by chimpanzees
all,
exposed
past)
by these
presented
from
were
recent
home range
represented
except
or
the
on
3
1960).
Tappen
Mt. Assirik
have been less
would
in
1972,
Above
reported
Rhine
and
and her
7 deaths
recorded
leopards,
by
a lion
car,
on baboons
predations
influence
Jolly
described
female
an adult
evolutionary
baboon's
dogs
of
chimpanzees.
(three
the
lions
(1970)
Altmann
(1980)
Rasmussen
a tourist's
due to
The baboons
predator
from
and Owens (1972)
all
mostly
of
case
possible
baboons
(1963)
Cilliers
one
and
of
being
saw baboons
Altmann
by a hyaena.
24 attempted
same period,
kill
probable
begging
to
described
leopard.
a
night.
been
"loss
the
one
at
had
that
by
one more,
by leopards
kills
(1973)
Nagel
infant
Haddow (1952)
1967).
and Turner
not treat
and there
mean troop
apparently
sizes
decreased
decreasing
with
3.12:
risk
of predation.
(DeVore
that
troop
extreme,
(Hall
stress
by
of
is
to
clumped,
the
to
one
food
food
allows
own
is
the
food
not
is
competition,
themselves
all
are
these
from
emerge
views
opposed
which
and feeding
renew
can
resources
Two
important.
certainly
predation
its
on
abundance
risk
At
1975).
The degree
size.
between
abundant
and Dunbar
food
that
which
with
other
the
severe
under
extreme,
is
beyond
1965).
Hall
and
sizes
There
size
entirely
(Dunbar
form
troop
for
account
speed
the
at
to
interaction
the
can
disintegrate
however,
likely,
sufficient
scarce
while
T. gelada
(DeVore
food
structure
1963a),
herds
It
abundance
troop
1970).
troop
troops
since
different
influence
does
and the
troops
especially
and Altmann
Altmann
in
review
unclear,
can have widely
home ranges
limitation
presumable
is
baboon
savanna
of
size
habitat
the
1965,
and Hall
evidence
large
in
overlapping
exploiting
and
food
of
abundance
the
between
The relationship
some
Introduction
Food and Size of Group:
considerations.
Firstly,
clump,
Lamprecht
others
especially
in
is
which
whole
is,
not
Post
This
so they
might
is
(1980)
well
denied
Subordinate
should
benefit
from
by
at
animals
forming
3.25
groups
work
a time
to,
access
animals,
first
field
of
of
enough
gain
access
to
food
by both
or
being
(1977)
Dittus
subordinate
from,
displaced
most
the
primates
captive
of
starvation
their
groups,
the
of
leave
then
food,
for
suffering
to
1975,
food
animals
studies
Wolf
by large
exploited
in
defend
each individual
a source
contain
are
accepted
support
being
best
showed that
who showed that
be the
is
there
as
find
on or
and
if
that
suggest
they
should
clumped
further
first,
sources.
competition,
them
clumps
et al.
and Wrangham (1981),
food
(1974)
(Caraco
lions
for
can
is
size
group
feed
of
variety
a wide
optimum
that
animals
example,
strongly
and has been given
animals
of
number
the
that
unlikely
for
and joins
order.
died
the
from
selectively
Thompson et al.
when such
group.
rank
it
1978).
monitors
food
by
as
is
it
sources,
directly
governed
one
since
food
plant
feed
baboons
group;
from
feeding
having
(Anderson
own
done
1980b,
Chepko-Sade and Sade 1979,
fragmentation
such
occur
(1974)
and Baker et al.
food,
foraging
abundant
individual
doing
of
In
competition.
should
seasonally,
in
is
food
is
that
can return
permit
more
(1980)
there
successive
resources
to
harvest
a food
to
Cody (1971,1974)
1971)
there
Pulliam's
argument
Watt's
and
suggested
the
if
idea
reduce
was
in
subsequent
the
animals
of
resources
Evidence
efficient.
swath
which
Small
groups
above-ground
has
that
should
through
the
area.
therefore
been
primates
grazed
be
productivity.
3.26
found
and
detected
is
in
are
capable
provided
areas
(1974)
at
both
that
less
foraging
of
by
slowly
would
detecting
by Wrangham
or
Hamilton
suggested
they
making
of
and avoided
They
area,
so that
1974)
Cody
as individuals
entire
grazing
previously
(1973)
(Cody
birds
form
A modified
which
the
over
(Cody
consequence"
a swath
foraged
between
time
the
between
is
passes
other
food
was scarce.
where
areas
Dunbar
exhausts
or
did not form flocks
finches
be applied.
supported;
a group
same number
of
Wrangham
Newever,
the
will
group
troops
by
the
than
the
by other
when
idea
rapidly
of
area.
competition
predators
not
could
that
group
the
were
which
way.
in
flock
feeding
"no
this
in
a similar
regulating
wintering
but did
little
was
and there
in
source
showed that
where food was abundant,
However,
advantage
that
be largest
fragmentation
to
grounds,
and
will
more
shared
are
vary
should
low food abundance.
expressed
the
feeding
competition
competition
themselves
of
patches
be little
might
visits
if
feeding
of
groups
(1970)
then
them,
increased
of
of
the
to
risk
on theoretical
predators,
renew
the
and group size
feeding
food.
of
at times
that
reduced
at times
both
exploitation
that
out
points
species,
if
efficient
groups,
and Watt
Hamilton
felt
that
expect
abundance
reduced
suggested,
for
watching
scarce.
suggesting
group
in
spent
(1981)
of
smallest
is
there
which
low
of
expense
times
(1976)
We should
groups
the
of larger
Pulliam
among animals
in
at
being
groups
with
or
then,
formation
By contrast,
time
badly,
general,
the
allow
times
should
Both Thompson et al.
reasonably
1978).
Rubenstein
times
a
(1976).
of
high
Food and Size of Group:
3.13:
Results
with
varied
species
the median count
thus
Stands
of
feeding
probable
is discussed
m2
nearly
probably
yr-1
all
its
at
by another
of
seen.
competition
This
observation
site.
productivity
of
least
(Chapter
this
but
2),
abundant
since
in
the
the
dry
wet
was probably
in
fell
no rain
was in
production
Mt. Assirik
at
season.
(other
season
Species
mode median
n
min
max
3
14
3=4
1.0
6
1
17
1
4.6
111
Afzelia
africana
Alophylus africanus
Anona senegalensis
1
2
1
14
2
2
1
1=2
2.3
1.5
17
1
6
Bombax costatum
1
1
1
1.0
2
1
1
4
30
2
1
1
1.6
1.0
9
21
1
dudgeons
digitata
Borassus aethiopum
Borassus aethiopum
Cassia sieberiana
in
below
2
1
8
18
18
1
20
1
5
Cordyla
pinnata
Diospyros
mespiliformis
3
1
3
14
Erythrophloem
suaveolens
Ficus spp.
Gardenia
erubescens
1
2
1
Grewia lasiodiscus
Hexalobus
Cola cordifolia
Cola sp. unknown
Combretum
Cordia
spp.
myxa
monopetalus
Kaempferia
aethiopica
Khaya senegalensis
-
-
2
-
2.0
7
-
1
1
1.0
14
1=5
3.0
2
2
3.0
1
6
20
17
1
1=2=4
2
4.5
4.0
12
13
1
1
3
1=2=3
2.0
3
1
5
1
1.0
4
1
1
1
13
1
1=6
1.0
6.0
5
-
-
the
5
dry
Food was
methods
Feeding Group Size
Adansonia
from
Feeding
Table 3.10:
Acacia
was
enough to
displacements
food abundance are used in Chapter 7).
determining
troop
than
any species.
be large
to
feeding
every
almost
plant
Numerous
were
less
An entire
unlikely
season.
either
was always
together.
were very
in
but
from a single
together
at
3.10)
travelling
above-ground
annual
(Table
of any
in Chapter 7.
further
1 kg
season,
feed
one animal
of
was therefore
around
animals
troop
whole
sites
The
species
any one species
the
support
of
to
able
never
the
in any one plant
seen feeding
The median number of animals
of
Species
mode median
min
max
1
3
1=3
2.0
2
Lannea acida
Lannea microcarpa
1
1
46
10
1
1
2.5
2.5
48
6
Manilkara
multinervis
2
2
Meliaceae
sp.
1
1
1
-
1
latifolia
1
4
Oncoba spinosa
1
Oxytenanthera
heudelotti
Landolp hia
-
n
1.0
3
1
1.0
9
2
1
1.0
3
1
3
1
1.0
8
Parkia
biglobosa
Pericopsis
laxifolia
Piliostigum
thonningii
1
1
1
5
4
10
2
1
1
2.0
1.0
1.0
5
4
6
Pterocarpus
1
17
1
3.6
32
1
1
Saba senegalensis
Sorindea juglandifolia
Spondias mombin
Sterculia
setigera
Strychnos
spinosa
1
2
1
7
1
18
2
2
7
1
Tamarindus
1
1
1
Nauclea
Raphia
erinaceous
sudanica
Trichilia
Urtica
Vitex
aby'ssinica
indica
prieuriana
sp. unknown
madiensis
Zizyphus
spinachristi
-
-
1
2.6
1
1
35
1
1.0
5
1.0
1
6
-
-
1
1
1
1.0
3
1
1
1
4
1
1
1.0
1.0
5
1
5
1=2=3=5
2.5
4
1
135
1
4.0
11
1
6
grass
Cissus spp.
sp. unknown (1)
sp. unknown (2)
1
5
1
2.5
8
1
4
1
4
1
1.0
1
15
-
sp. unknown (3)
-
4
1
3.14:
Food and Size
Feeding
in prep,
or implied
1965a, Hamilton
(1976)
Pulliam's
reduced
study
nor
suffer
and risk
by observations
feeding
troops
et al.
baboons and other
studies
has
1968b, Dittus
1978 Homewood 1976, Ransom 1971, Smuts
that
form large
should
page 3.3);
on mixed-sex
nor
was it
herds
(Wrangham
of
1976),
3.28
both
suffering
animals
was not supported
competition
(Chalmers
primates
1979, Wrangham 1976,1981).
of predation
(see
of
in many field
view
food availability
on P. papio
Discussion
and Leland
Struhsaker
competition
in
competition
been observed
1977 Hall
Group:
of
8
4
5
groups
by observations
in
Homewood's
T. gelada,
feeding
in times
of
made in this
(1976)
study;
whose members may
and whose adult
males
suffer
threat
the
"predation"
of
herds
animals,
larger
where
are
feeding
competition
times
In
Discussion
determining
example,
(Steward
1955),
(Silberbauer
forage
to
related
to
the
the
causes
1972,
the
Duncan
Vigne
food
Maus 1906),
the
and the
concentrated
at
prey
(eg
available
(eg
water
to
unrelated
Woodburn
ecology
may
be
1975)
or
may have
1972),
or
the
Birhor,
(eg
to
human
other
groups
of
Lee
! Kung.
the
In
Hadza,
the
1937)
dispersed
and
aggregation
and
bushmen
(Warner
year.
of
group
Indians
G/wi
the
sources,
times
other
at
dispersal
of
food
in
Shoshonean
Australia
of
little
or concentration.
abundance
Northern
that
1979),
at
all
flies
tsetse
times
seems probable
at Mt. Assirik
baboons
of
since
the
in
changes
groups
of
sizes
abundant
(page 3.3)
of predation
was largest
changes
seasonal
Murugin
of
In summary, it
size
conditions
plays
apparently
1962),
unlikely
influenced
erratically
for
Sinha
1974).
seems
and
in
changes
resources
size
group
However,
(Damas 1972,
apparently
Williams
It
At Mt.
evidence
size
(Meggitt
availability
social
and
Changes in Group Size
on the
size.
to
seasonal
the
group
Aborigines
seasonally
the
that
1966)
(1976)
Pulliam's
baboons,
these
1973).
was
Walbiri
on scattered
societies
(Dunbar
As there
have predicted
group
1972),
congregated
(Crook
food
scarce
predation.
predation
Eskimo
the
more extensive
Causes of
of
be related
can often
size
are
of
In
1976).
abundance.
human societies,
in
part
For
low food
of
times
the
amongst
(Wrangham
groups
the
of
and we should
General
3.15:
grass
of
was some risk
were satisfied,
in
in
smaller
swards
there
Assirik
by all-male
of
(as
it
may do
(Glossina
horses,
in
morsitans)
were
year.
that
seasonal
in
baboon
group
changes
came about as a result
at a time of abundant
of
abundance
flies
biting
food,
of low food availability.
3.29
of
and a decreased
increased
risk
risk
at a time
.,
.`
CHAPTER4: Activity
Budgets
INTRODUCTION
4.1: Ecological
Different
in
example,
an increase
different
in
results
(Nagel
in
some loss
time
1973, Oliver
spend
less
time
spend
less
time
spent
(Schoenen
in
a "currency"
food
1971),
or
shortage,
studies
time
more
spend
in prep).
and Dunbar
and Lee 1978, Iwamoto
at
day which
of
Several
food.
baboons
For
by feeding
a time
at
than
other
activity
conditions.
ecological
may be achieved
feeding
of
of
to
their
balance
apparently
response
day
in times
have shown that
in
ways
times
profitable
baboons
of
groups
budgets
less
on Activity
Constraints
feeding
and either
1981),
Post
1978,
(Oliver
Lee
or
behaviour
in social
and
Nagel
in
Dunbar
(Dunbar
Iwamoto
1980,
prep:,
and
resting
1973).
in food availability
Seasonal differences
than they do between neighbouring
troops
The time of day at which various
environmental
frugivorous
frugivorous
animals
larger
a social
but
group
the
larger
its
within
the social
it.
with
with
about 250 members, there
group
each
exchange, and hence the troop
proportion
The increase
has
to
may spend a greater
u. 1
to
may lead
its
greater
of the day given over
the possible
is rapid;
for
dyads, while
longer
spend
proportion
Anecdote's
for
Fable's
In a larger
dyads.
31,000
over
are
probably
of
The number of
group the greater
are 9,045 possible
135 members, there
individual
companions.
competitors
of
only
not
the time budget; for example in a
influence
number
However, the larger
consists
social
food, and a greater
in finding
number of relationships
social
at
(eg
than competitors
earlier
environment
also
in the group might well
to feeding.
troop,
the
animal,
surroundings
difficulty
troop,
to arrive
birds).
For
physical
the largely
intense,
is
7) might attempt
trees
food sources such as fruiting
renewing
are seen may change with
competition
(chapter
baboons of Mt. Assirik
of neighbouring
(Homewood 1976, Nagel 1973).
activities
When feeding
changes.
home
more between seasons
to differ
budgets
which may cause activity
ranges,
within
between the habitats
than the differences
be greater
one home range may
in
social
of the day in
interaction,
social
perhaps at the expense of resting
Coalitions
individual's
(Varley
competitive
de Waal
agonistic
encounter
can
on
rely
the
is
encounters
are,
in
than
more
typically
troop
and
very
30,000
However,
we might
between
juveniles
presumably
baboons,
large
spend much time
Seyfarth
Mt.
at
1980 and Wrangham
larger
troops
smaller
troops.
troop
but
in
support
1981).
more
time
for
one
grooming
of
in
predict
another
this.
but
also
more
1980).
collecting
including
monkeys,
other
therefore
on
formed
be
coalitions
also
1975, Dunbar
1976,
in
1966,
Symmes
and reviews
and
We should
Datta
males,
suitable
and Dunbar
(Dunbar
in
(Rasmussen
groups
each
females
effect
to
species
several
one another
can be involved
among adult
were
cannot.
are dyadic
there
little
not
agonistic
(review
1979),
an
that
Adult
matrilines
small
Assirik
1976, Varley
spend
in
regularly
grooming
1980, Strum
Fable's
Watanabe
than
groups
which
than
especially
of
that
rank
equal
have
and
in
baboons
associations.
would
1977,
on coalitions,
animals
of
of
Varley
Any individual
triadic
their
coalitions,
(Massey
conditions
directly
three-way
size
Observation
data
each member of
different
expect
one
many more possible
members
in
other
than
other
frequently
or
do better
each group;
support
1981),
to
1980).
food
outcome
probable
Seyfarth
more
1977,
an
over
conflicts
(Cheney
the
may alter
another
in
a rival
reversing
one
of
against
partners
(Kummer 1968b,
of course,
in
sexual
and in
support
animals
against
or
1977)
likely
relationships
more)
ability
Symmes 1966)
and
Symmes 1966,
There
two (or
between
time.
animals
in
do animals
in
that
than
Goosen
METHODS
4.2: Activity
Budgets
budgets
Activity
social
activity
animals
the animals
easy to
relatively
observe.
were recorded
categories
were determined
included:
often
At
sat
these
at intervals
sit,
stand,
in
open areas
times
activities
of 20 minutes.
walk,
4.2
(1) During
in two ways.
eat,
of
in which they were
of
The principal
self-groom,
times
all
visible
activity
and watch observer
(all
activities
during
with
any other
baboon),
Activities
chase.
because active
space
sedentary
scan
time,
of
recorded
together
usually
interrupted
animals
to
the
purposes
one
four
This
with
activity
from
that
is
problem
cumulated
across
of
the
active
Kolmogorov-Smirnov
same troop
across
of
of
differential
seasons
selection
of
summed within
two
troops
of
records
two-sample
the same season.
u.3
for
test,
the
with
the
same activity
to
any
different
and
by
into
each
of
both
for
each
each
in
(1972)
compared
concerning
troop
of
falling
then
so
subject
seasons
were
day and results
and with
not
by Clutton-Brock
were
from
combined
either
visibility
observations
activity
was extracted
within
is
For
the activity
follows
part-day
represented
discussed
the
proportion
from
data
number
categories
budgets
the
(1972)
the
hour,
on the
my
(1979).
my notes
The data
hour.
records
method
The total
The
the
after
contact.
was
this
Clutton-Brock
from
was no record
day was equally
This
apart
activity
record
the
of
squared).
the
first
season.
bias
or
there
together
Homewood (1976).
using
if
in the
activities.
The
hour,
visible
In
and Leland
of Struhsaker
extracted
the
behaviour
of
I
moving),
their
of
each
were
animals
or
caused
assessment
towards
My appearance
made.
frequently
of
the
saw
a
and regular
feeding,
was
sampling
sampling
a
within
animals
observe,
resting,
seconds
I
with
biassed
the
I
whenever
and
instantaneous
analysis
of
each hour
obvious
the
the
this
follows,
all-day
few
first
the
durational
on the
mentioned
any
the
than
recorded
that
in
together
When
to
record
so that
sight
of
resembled
sampling
behaviour
and
mount, copulate,
been
difficult
the
both
analysis
Focal animal sampling was
(2)
(social,
which
at
their
move out
made
rather
time
the
recorded,
have
would
Instead,
activity
interacting
not
tended to move out of sight
usually
impossible.
the
(in
play,
necessary.
grooming.
as
were
predominant
with
normally
they
were
samples
the
if
sampling
such
feeding
or
suckle,
were also
animals
and
activities,
foraging
list
of the behaviour
not attempted
short
under "groom"),
not on this
description
full
was apparently
animal
groom and be groomed
are included
categories
the
which
troops.
(using
were
activity
activity
compared,
same activity
across
chi
troops
in
in
RESULTS
4.3: Activity
Budgets
left
The baboons
06: 46
between
or in
ravine
After
the
for
ground
At
troop
moved off
chi
earlier
as it
whereupon
behind
food
wholly
socialise,
would
they
before
trees
in
the
dry
to
it
or
while
in
part
when
reach
spent
At some time
areas.
a sleeping
the
the baboons
by nightfall
climbed
There
night.
into
and were
the
never
the night.
4.4
known to
area of
to
less
the
day,
lagging
to
and
rest
troop
one at
which
behaviour
were always
the
might
the
some social
The baboons
descend
or
apparently
afternoon
the
test;
troop
across
one from
was often
the trees.
more
sometimes
late
day's
its
(medians
The scattered
day,
might
some animals
with
the
07: 46 and 08: 00)
stayed
foraging
mid to
a different
often
site,
previous
in
forming
however,
began
between
troop
on
interact
troop-mobilising
together
moving
dozing
behaviour
Usually,
time
out,
the
throughout
the
a pseudopod
The troop
moved on.
others
bordering
to
08: 16 and 08: 30)
and spread
= 1;
sat
often
with
the first
entered
df
starting
cases
The
p<0.02).
time
troop-mobilising
sight.
between
(median
in
social
and engaged
they
seen,
(median
time
fragment
sources
plateaux
gradually
these
time
= 9.59;
squared
progression.
in
season
season
trees,
period
observer's
= 1;
occasionally
unproductive
social
although
of
df
before
a general
out
began
the dormitory
(median
season
wet
chi
was sometimes
moved, until
it
at
the
into
the
laterite
on
more
the
= 7.06;
in
in the wet season
in the wet (median
squared
together
or
abruptly
place
did
reform
an
gradually
changed
it
hour
dry
the
test;
either
nights
stormy
did
they
beneath
end of
1979)
Stolba
which
movement
sat,
in
earlier
(medians
the
socially.
(Kummer 1968a,
have taken
then
the valley
behaviour.
than
07: 00)
and
They
p<0.005).
than
06: 45)
06: 31 and
between
trees
the
ground
in
the
during
Figure
4.1:
The estimated
mean number
in various
age-sex
classes
both seasons.
Adult
80
per day
during
of minutes
activities
males
Wet Season
Dry Season
8
aý
Fable's
40
h
Troop
Anecdote's
Troop
W
?
0
W
r'
C
by different
times
in
spent
social
3
=
cß
Q1
mcý
O
ö
s
u
a
w
N
120-
W
pQa
ar
ä
1W
Adult
cq
1o
.t0
females
Wet Season
Dry Season
ti
C)
a
N
C)
Fable's
Troop
Anecdote's
Troop
C
Cr
J
C
cl
ý
Q)
cd
O
D
Z
h
A 60
ý
I
Juveniles
U,
cr.
gW
"N
E'+
R
Season
Infants
8
cz,116V
ý
yý
8'
mW
19
ti`
C)
Wet
Dry Season
ti
E 30
c"
ý ow
J
w
J
w
_
3c
''`
wä
i
=
z0
LLJ
tý
4.3.2: Distribution
of Sample
Each hour
times
number of
troops
day between 07: 00 and 17: 00 was sampled the
of the
within
but
seasons,
the
sample size
same
between
differed
06:
00
18:
00
fewer
of,
and
were
a
sampled
-.
Table 4.1 summarises these sample sizes.
The hours
and seasons.
number of times.
Table 4.1:
(number of samples of each hour)
Sample size
Troop
Season
1
06: 00
07: 00-17: 00
Fable
Wet
Dry
I
I
12
0
253
143
Anecdote
i
Wet
4.3.3:
Absolute
Frequency
Both troops
in
the
troop;
dry
20
2
187
20
(chi
No seasonal
squared = 6.02;
df = 1;
differences
(Table
time spent resting
In
I
I
I
274
177
I
262
I
210
Activities
of
squared = 31.05;
chi
220
were seen to feed less
season
14
26
20
I
Dry
18: 00
in the wet season than they were
df = 1;
p<0.02
for
p<0.001
for
Fable's
troop).
Anecdote's
the time spent moving or for
were found for
4.2).
Table 4.2:
Activities
the day
Frequency
of
of Observation
throughout
of Two troops
Troop
Season
Activity
Moving
Social
Feeding
Resting
Total
Fable
Wet
Dry
102
80
68
22
42
47
62
28
274
177
Anecdote
Wet
93
56
49
'64
262
27
81
36
210
Dry
No differences
data
for
the
66
were found
two
troops
between
were
the
pooled
u. 5
two troops
for
the
within
comparison
seasons,
of
and
activity
budgets
in the wet and dry seasons.
Baboons at Mt. Assirik
engaged in
in the wet season than they did
df = 1;
season than in the wet (chi
A total
the
on Fable's
(1102 in
troop
the dry
p<0.005).
were made in
1841 were collected
of which
(655 in
troop
in
activities
the wet and 739 in
on Anecdote's
were collected
df = 1;
of individual
behaviour,
social
squared = 11.38;
more frequently
squared = 22.84;
of 3093 observations
scan samples during
more frequently
in the dry season (chi
and were seen feeding
p<0.005),
behaviour
social
the dry
and 1252
season)
the wet and 619 in
dry
the
season).
The most
which
common activity
44.0%
occupied
passive
run,
animals
recorded
eat,
groomed other
< 0.05).
adult
similar
age-sex
class
its
time
age-sex
various
4.1)
spent
the
did
those
the
of
of
from the
animals
(chi
troop
Anecdote's
males from both groups
adult
(chi
frequencies
squared = 0.02;
from
on
the
(chi
activities
p always
< 0.025).
among the
squared,
of
of the
adult
and juvenile
grooming
in
the
p<0.01
in
each case)
day
and almost
4.6
all
in
by
between
different
class
age-sex
in
spent
baboons spent
did
spent
those
from
< 0.025).
The results
wet season as they
pooled
time
activities
are
shown in
twice
about
the dry
age-sex
activity
social
per day spent
number of minutes
was calculated.
of
Each
various
squared,
therefore
were
was
p always
the
(chi
seasons
The proportion
various
known proportion
activities
within
two troops
the
seasons.
differently
classes
troops
by each age-sex
activities
on various
two
within
an estimate
In both troops
time
squared,
From the
(Table
as
walk,
49.4%
occupied
groomed other
In particular,
across
classes
(chi
distributed
other
troop
than
The results
age-sex
seasons
of
of
< 0.05).
p never
each
(sit,
baboons
other
or
active
as
ns).
was
within
either
observer)
males at equivalent
The proportion
class
of Fable's
category
p never
animals
was grooming,
scans.
more frequently
same category
df = 1;
the
watch
times
social
involving
not
self-groom,
in
No age-sex
squared,
scanned
Activities
partners.
stand,
the
of
during
recorded
Fig
classes
4.1.
as much time
(two-tailed
in
on the
both
t test,
troops
O
O
N
r
ti
I-
cl
A
Cl.
G
o ý-
ow
o
N
oC
N
U
E
E-
0
0
N
G
00
CD
C)
It,
o
sp, xoýaý
qu-
Jo
31100,zad
00
W
c;
G)
0
sp. zoaag 20 . ua3Jad
8
N
c"
4)
C) Q
hiL
"0
a,
c3
r
0
ow
00
o0
0
N
r
C)
E-
%b
00
CD
In
0
0
rQ
S
0
O
O
40
In'
w
0
4.2:
Figure
Probability
in
of finding
members of two troops
times
at various
of day.
A Percentage
in which Fable's
of hourly
records
feeding
in two seasons
or moving
B
C
D
Percentage
of hourly
feeding
or moving in
,
Percentage
in
engaged
Percentage
in
engaged
Adult
Adult
males
females
in which. Anecdote's
records
two seasons
of hourly
record
behaviour
social
of hourly
records
behaviour
social
behaviour
social
troop
were
troop
were
in which
Fable's
troop
were
in two seasons
in which Anecdote's
troop
were
in two seasons
spent more time
in
dry
the
in
no difference
p always < 0.01).
test,
Anecdote's
in
the
troop,
There
spent
may have
order
time
of
median
group
4.3.4:
Distribution
The
been
for
size
two
the
feeding
or
in
to
at
a similar
seasons,
troops
and within
The probability
however,
a similar
probability
extremely
low
wet
season
(Fig
were
of
behaviour
occupied
afternoon,
when,
17: 00.
seen
In
moving
11: 00 or
or
12: 00,
feeding
less
in
from
in
In
of
day
the
differed
about
the dry
60%
about
of
summary,
time
climbed
to
80-90% of
season,
08: 00,
dry
records
until
records
by 08: 00,
remained
25% by
about
likely
to
trees
troops
feeding
the principal
ones,
or
were of
feeding
were either
and in over
about
until
were of
be
90% by 10: 00,
accounting
for
18: 00.
behaviour
with
season both
late
to
mid
about
the
social
season
the
the
was
10: 00 in
increasingly
left
and by
behaviour
after
the
records
they
dry
the
social
until
were
stayed
about
records
in
behaviour
of the day between 50 and 90% of records
which these activities
In
it
troops
the
time
which
over 50% of records
times
season
feeding
at
troop,
both
of
while
10% of
than
Fable's
wet
or moving.
or moving
behaviour,
in
within
pooled.
In
until
or
seasons
troops
across
season.
two
the
social
troop
10-20%
About
social
For the rest
moving.
after
the
4.2).
in
likely
either
in
day
therefore
were
wet
behaviour,
social
Results
was only
finding
of
in
of
troop
the
of
order
rank
the day
seasons,
in
the
since
group
the
time
of
amount
of
same as
time
either
10: 00
and
two seasons.
ns).
across
probability
10: 00 the
the
the
size
be engaged
test,
finding
of
30% until
about
between
across
likely
2-sample
There was
by juveniles
play
and the
two troops
resting
was no statistical
p always > 0.05).
test,
was
were
in
spent
grooming
(Kolmogorov-Smirnov
above
time
of Activities
troops
was for
in the wet season.
sitting
and juveniles
spent
The exception
whom there
some correlation
by adults
grooming
moving,
for
amount of
in the two seasons (t
infants
in the wet season than they did
nothing"
in the amount of time
difference
rank
(t
season
males
adult
"doing
sitting
the
the
characterising
season.
4.7
In
the
wet
troops
season
at
different
the
baboons
socialised
until
and began to
mid-morning
day.
In the dry season social
before
09: 00.
4.3.5:
Comparison With Other Troops
One other
budget
activity
different
more
activity
resting
to
unable
biassed
Table
profile
less
and
follow
the
towards
and Nathan
P. papio
of
in
the
wet
from
that
in
baboons
day,
all
the
provides
data
on the
Their
data
show a
season.
this
study
they
so that
(Table
4.3),
with
that
they
were
state
their
budget
activity
is
4.3:
Activity
of
Budgets
P. papio.
Group
Dunbar and
Nathan (1972)
40.2%
20.5%
19.2%
20.2%
Anecdote's
troop
33.7%
17.41%
27.8%
21.1%
Mt. Assirik
36.9%
18.9%
23.5%
20.7%
from
troop
given
any
from
of P. anubis
populations
of
than did
in
species
troops
2 populations
their
dry
they
those
time
habitats
spent
(t
in
the
dry
moving
(Aldrich-Blake
In
species.
in social
4.10;
=
(Harding
4.8
baboon may manage their
other
of T. gelada
more time
troops.
of
of
spent more time
P. cynocephalus
spent
of
Resting
46.6%
populations
although
of
26.1%
differently
P. anubis
Populations
15.8%
Troops
P.
two
9.1%
Fable's
rather
of
Activity
Social
Feeding
Moving
either
in
when the baboons were not moving.
periods
Comparison
time
1972)
However,
moving.
later
gave way to moving and feeding
activity
(Dunbar
study
move and feed
(t
and less
et al.
but
less
df = 7; p<0.025).
= 3.01;
Dunbar
than did 3
p<0.005)
like
season rather
1976,
7
particular,
behaviour
df = 8;
time
the
troops
and Dunbar
time
feeding
of
1974c)ß
than
did
1971, Dunbar and Dunbar
1974c,
Hall
1965b,
Hamilton
Nagel 1973, Oliver
prep,
al.
1975,
1978,
et al.
Iwamoto
and Dunbar
in
pers comm, Rasmussen pers comm, Van Citters
et
1967).
DISCUSSION
4.4:
Activity
The
Budget
(Post
budgets
activity
qualitatively
to
similar
Period
baboons
the
of
those
P. cynocephalus
of
troops
within
more in
the
budgets
as
two troops
spent more time feeding
In
the
of
wet
of
Anecdote's
trend
and accords
frequencies,
common in
than
in
the
so that
moving
or
or
(82
min),
which
goes
and
infants
the
dry,
made throughout
foraging
more in
would have been introduced
the
probably
the
day,
went
the results.
4.9
are
other
data
play
and
possible
troops
the
both
constraints
less
groomed
the
basis
the
of
may have
not
been
time each
than
more
This
predicted
grooming
no
in
the
observation
more
bouts
in
may
the baboons
while
If
such
the dry,
bias
undetected.
did
wet
Since
were collected.
it
did
troop.
play
largely
the wet season than
into
maintain
in Anecdote's
did
which
to
against
to prediction.
contrary
order
the
to spend in grooming.
on
classes
apparently
the manner in
occurred
that,
were
larger
being
grooming
they
in
energetic
afford
mins)
age-sex
animals
they
spent a little
probably
observation
than
in
in
within
troop
to
were not
behaviour
with
Fable's
be due partly
were
group
coalitions
Juveniles
season
(77
season
those
troop
in
particular,
in the dry season,
males in Fable's
did
than
there
although
groomed each other
that
suggest
may have reduced the time the baboons could
Adult
large,
each other
behaviour.
this
of
were
grooming
bonds,
social
troops
may
but differed
were similar,
and juveniles
This
predicted.
interpretations
scans
were
in Kenya
in Amboseli
Adults
when the
spend more time
strengthen
the
of
between seasons.
wet season,
season,
groups
day
Assirik
Mt.
at
1981).
The activity
dry
Social
within
About
half
apparently
might
in
solitary
dozing
either
activities
or
watching
study,
especially
in
socialising
continuously,
in
since
"resting"
a
the
with
the
I
other.
since
times
collected
largely
throughout
more
the
(1973)
than
26% of
suggests
environment
of
their
that
the
animals
the
that
time
(about
combination
in
the
16-17%)
of
dry
X1.10
in
dry
baboons.
resting
habitat
season
and
than
and
to keep the
in
social
possibly
"social"
not
periods
periods.
he
observed
the
season,
Baboons
in
varied
category
one
all
baboons
the
and
probably
and were
"rest"
of
the
in
the
morning
not
since
resting
behaviour
of
baboons
the
resting
in
It
and "resting"
was more prudent
early
certainly
overtax
group
sitting,
observer.
animals
first
it
animal
the
of
frequencies
of
time
not
that
in
activities
their
more taxing
of
on
suggested
did
marginal,
less
the
"social"
data
day and almost
Nagel
though
spent
of
or
however
felt
separate
representative
proportions
were
periods
"social"
categories
the
majority
categories
were
the
the
including
animals,
separate
"social"
involving
often
other
to
during
recorded
activities,
seem artificial
this
then
the
of
the
troop
was the
at
dry
habitat,
Mt. Assirik
season,
sizes
habitat
spent
which
made their
at
Awash.
CHAPTER5: Ranging
INTRODUCTION
Without
biomass
the
ecology
in
of
region.
The
by which
troops
the
of
determined
part
sites,
several
baboons
in
the
ranging
of
baboons.
This
NK, since
Dupuy
from
in
observation
the
(DeVore
malaria
1975,
File
Dunbar
Hausfater
1975, pers
their
exploit
obs)
respiratory
and Moore
be attempted
should
environment,
the home ranges
of the
and what governs
(Deag
1973).
meet
the
of
the
of
population
thought
known or
the
GPN de
the
to
the
control
of
ranging
relevant
to
suffer
(Nagel
mange
1973),
Douglas-Hamilton
1965a,
parasites
1973, File
their
overlap.
each might
in
(Dunbar
1980a,
(Dunbar
1980a,
Finally,
prep).
how the
we understand
until
part
infections
and intestinal
1976, Kuntz
and Watson
no management
McGrew),
one or
might
(DeVore
share
which
of
infectious
including:
schistosomiasis
comm to
and Dunbar
are
is
an area
have any means
are
troops
especially
plagues
in
yet
park
knowledge
baboons
diseases
1965a),
pers
that
Certainly,
many infectious
the
troops
perhaps
remarks
park.
in
to have on
ranges,
We do not
its
estimate
likely
baboons
known how large
which
is
overlapping
members.
include
is
it
of
how many other
must
area
(1971)
have
we cannot
that
pool
baboons
determine
the
gene
yet
or with
patterns
epidemiology
baboons
we do not
are
park
the
or not
whether
demes because
Since
influence
exchange
and which
estimating
area of a troop
area and hence the
per unit
sleeping
the home range
knowing
animals
movements.
5.2: Size of Home Range
The size of an animal's
(Brown 1975, Hunter
foliverous,
Milton
size,
the
view)
or
and May 1976).
terrestrial,
same order
(Clutton-Brock
(Clutton-Brock
Thus all
troops
and omnivorous,
of
et
upon the degree to which it
and partly
omnivorous
see Turner
magnitude,
depending
5.1
rate
al.
(Harvey
1970 for
a
is frugivorous,
1977, McNab 1963,
of Papio baboons, being similar
should
and Harvey 1977,1979,
and Harvey
body weight
on its
and May 1976) or metabolic
1964, Milton
1981, McNab 1963, but
and Clutton-Brock
conflicting
home range depends in part
in
have home range areas of roughly
partly
on group
DeVore and Hall
size
or weight
1965, Suzuki
1979).
(Bernstein
densities
local
the social
space
time
of
essential
water
et
1976,
al.
Saayman 1970),
1972,
Clutton-Brock
Stoltz
(Altmann
by the
1974,
1970,
DeVore
and Orians
Brown
1971,
Hion
Stoltz
1973,
and
1970, Kummer 1968a)
1973,
and Buetenandt
in
distribution
Gautier
and Keith
and Altmann
1977.1979,
1967b, van Winkel
Altmann
and
1976,
Casimir
and Harvey
Struhsaker
resources
(Altmann
Harding
sites
and shape of home ranges of many
influenced
are
(Altmann
sleeping
(Bernstein
food
1972, McNab 1963, Nladik
all
baboons,
including
Hamilton
as the size
as well
and certainly
and
1970),
upon
and May 1976).
The location,
primates,
1965), and possibly
(Bernstein
of the troops
organisation
1975, Milton
1972, DeVore and Hall
1974a,
Clutton-Brock
and Hall
1975,
Hiadik
1965,
and
1975).
5.3: Movements of Baboons
(1979)
Rasmussen
behaviour
and hence social
animals
to
are
well
the
daily
decided
movements
at
that
the
day's
ranging
by the
(1979)
Stolba
the
baboons
and Sigg
one
group
of
the
day's
at
food
(Harding
considering
how day range
with
independent
variables,
to sleeping
sites,
length,
they
including
land
form,
spread
plan
(in
press)
(in
habitat,
largely
demonstrated
movements
during
the
has been shown to be affected
troops
could
size,
were
press)
season,
be
humans, and by
or
these
speed of movement of the
group
that
demonstrated
complements
areas
of
daily
its
control
baboons
their
other
chapter
entered
social
affect
and we now understand
and
Byrne
and of
This
1978),
P. hamadryas
of
Troop
1976).
which
and Stolba
and water
visibility
might
coordinated
by vocalisation.
of
in Baker
ranging.
Mt. Assirik
use
range
The movements of many species
troop
of
start
presence
frequency
(review
how a baboon
of
patterns
organisation.
researched
some extent
movements.
that
showed
time
and the
troops,
related
by
findings
to
several
of day, proximity
and climate.
5.3.1: Group Size
The length
of
the
daily
determined
by group weight
a species,
large
groups
tend
range
of
(Clutton-Brock
to have longer
5.2
is
partly
and Harvey 1977), so that
within
a group-living
day ranges,
animal
and therefore
move
faster
do smaller
than
on average
Dunbar in prep).
(with
troop
Fable's
1980b, Iwamoto and
250 members) should
hme-
have
therefore
than did Anecdote's
day ranges and/moved faster
had longer
(Anderson
groups
(with
troop
135
members).
5.3.2: Time of Day
Speed of movement is
Blake
1968a,
Sigg
(Chapter
the baboons
be
to
likely
be entered
earliest,
to contain
food.
first
of
There
slight
entry
would
presumably
fall
baboons
few
hours
the
of
they
tended
the
sometime
when
social
in
the
baboons
were
by
time
day;
of
daily
the
the
these
activity
or
likely
with
pattern
median
time
the
of
middle
to enter
should
habitats
into
so that
baboons,
of
patterns
to socialise
by entry
be unlikely
would
day,
in which
have been no consistent
to
the
first
should
The activity
influenced
at mid-morning
importance
Furthermore,
been
in which
habitats
followed
1979).
have
1974b, Kummer
Clutton-Brock
the habitat
that
also
should
those
1970,
Stolba
press,
4) suggest
found
that
specifically,
habitats
in
and Stolba
(Aldrichday
of
by the time
affected
and Altmann
1971, Altmann
al.
et
frequently
habitats
day.
in the
had
feeding
of
high
priority.
5.3.3: Season and Climate
1970), and frequency
in
low productivity
Times of
prep).
ranges
of entry
are
(Baldwin
longest
1978), others
Although
(Stoltz
do not, despite
ranging
air
that
some groups
strong
the distance
was apparently
day (Hall
seems to have little
related
1962a, Stoltz
effect
Struhsaker
which day
1967b,
1962a, Oliver
differences
and Lee
between seasons
in press).
moved by a troop
in the course of a day's
to the maximum temperature
reached by the
and Saayman 1970), although
temperature
on the speed of movement of baboon troops
5.3
van
of baboons show seasonal
1965, Hall
climatic
and Saayman 1970, Sigg and Stolba
In some studies
1972,
in
those
between season and day range length
in day range (DeVore and Hall
differences
sometimes
are
and Baldwin
Roosmalen 1980), but the relationship
in baboons is not clear.
1971, Waser 1976, Lee
(Lindburg
an area
into
and Saayman
1976, Stoltz
(Harding
speed of movement
Season affects
(eg
1976).
Harding
1981),
making
speed
of
For
it
when both
can
air
closely
related
the
an
wet
little,
the
57.6±7.6mm Hg at
16: 00 hrs.
through
day in
the
Since
closely
to
related
temperatures
the
the
cooling
the
enters
physical
sites
exert
Stoltz
(Sigg
Hion
of
any given
have
1971, Hall
in
ineffective
deficit
season
little
changed
have-been
it
than
the wet
but highly
exertion,
the
more
was in
the
dry
effect
of high
distance
roughly
then
might
from
the
are
19 sq km spent
be that
the
the
nearest
probability
proportional
(1976)
more than
of
to
5.u
the
to
model
probability
sleeping
of their
half
of
site
finding
inverse
them in
of
the
press,
living
waking
hours
account
for
being
found
their
fell
in
baboons
that
found
animals,
and Altmann
and Stolba
1972, Sigg
The simplest
site.
(Altmann
off
its
Sleeping
those
of
ranging
sites
sleeping
1979).
with
by
influenced
Stolba
press,
the
1965a, Rowell
frequency
moves and the
apparently
in
over
few
with Habitat
associated
both
Thus Harding
a sleeping
baboons
were true,
be
to
of movement might
baboons
of
influence
which
of over
200m of
ranging
07: 00 hrs
have ameliorated
and Stolba
and Saayman 1970).
within
would
areas
strong
baboons,
in a home range
this
certain
a
1970, Gautier
a troop
which
environment
including
deficit
pressure
in mid-afternoon.
The speed with
it
At
at
pressure
wet
would
(Moen 1973).
(10.5±2.4mm Hg), while
even slight
5.3. '$: Geography, Topography, and Variables
which
air
the
air,
is
9.9±3.8mm Hg at
from
to move
panting
or
mean vapour
16: 00 hrs
speed
in
temperature
when evaporative
of
vapour
to
cold
deficit
after
humidity
tend
near-saturated
was therefore
became moist
two.
Because near-saturated
sweating
varied
the wet season,
humans, might
by
at
Sweating
in the dry season.
effective
season,
it
season
when human skin
highest
the
of
with
itself
cool
at Mt. Assirik
season
dry
like
can
pressure
and was at its
end of
season,
can
to the vapour
the
interacted
were high.
than
more moisture
between
relationships
and combinations
temperature
and humidity
animal
the
humidity,
and baboons,
movement,
(McGrew et al.
are excellent
detail
in
examine
that
temperature
which
end of
varied
of
absorb
by
be
might
speed
amount
the
to
possible
it
example,
less
from Mt. Assirik
data
movement and temperature,
influence
hot
Climatic
linearly.
any given
square
the
at
If
quadrat
of
their
distance
from that
site.
The frequency
home
(Chalmers
al.
may be
range
1968a,
1977),
and find,
areas
speed
habitat
me to map the
frequency
of entry
land
determines
the
vegetation,
it
topography.
This
If
the
of
be
effects
on
exposure
to
the
of
and course
possible
speed
of
was therefore
movement
of
to rain
streams,
was compatible
with
from
patterns
in Chapter 2.
in the dry season when
season baboons should
In this
over 50°C.
affects
before.
primates
have been most pronounced
areas where this
and
and sun and
hence
and
ranging
predict
for
land
have tended to be found in more shaded areas and to move rapidly
shadeless
allowed
photographs
It
no
different
into
to the baboons, as I suggested
were frequently
As yet
area.
2).
speed
of the home range.
has not been attempted
should
sun temperatures
habitat
that
suggest
entry
Aerial
of
the
shade were important
then its
of
area
parts
runoff
of
may
frequency
or
more
behaviour
social
the
of
(Chapter
effect
affects
rate
Pyke et al.
evenly
the
in
time
for
preferred
productivity
time
their
more
field
different
into
form
spend
are
1975, Waser 1977).
distribute
have been attempted.
the
examine
but
1973).
movement
the habitat
to
Since
of
type
possible
which
the
to
do not
them,
those
related
of
with
to
1971, Nagel
et al.
be inversely
correlations
in
or
(eg Aldrich-Blake
will
baboons
that
vegetation
1976, Pyke et
1973, Post
(Glander
trees
the
their
of
parts
of
Nagel
fruiting
of
available
habitats
productive
distribution
the
1973,1975,
by that
in various
found
are
by
Clutton-Brock
habitats
the
over
animals
influenced
especially
Thus we expect,
which
with
the
through
needs of
other
the
troops.
In Chapter
3I
have led to a higher
baboons should
rapidly
through
understory
in
would probably
parts
of their
risk
that
them if
large
visual
of predation
have tended to avoid
they entered
density
in that
ranges.
in the wet season might
If
season.
this
them.
In the dry season, however, the
was burnt
off,
to encourage them to enter
In the dry season the troops
more frequently
poor visibility
them than they did areas of low visual
5.5
were true,
and to move
areas of poor visibility
areas of the home ranges
have found little
areas of relatively
through
suggested
density.
should
and baboons
such desolate
have entered
and moved more slowly
METHODS
5.4: Day Range Maps
The location
In many parts
possible.
of
photograph
its
of
habitat
position
In
points"
estimate
areas
the
on
In
from
three
the
counting
and 5 part
travelled.
high
was very
4.3km
results
on
the
(r
aerial
200m or
taken
take
on
any
bearings.
the
the
day
was
"half-hour
and these
baboons
continuously
were
I was unable to
between successive
points
line
formed the basis
by pacing
the baboons
at
for
1+.7km on
used here were taken
the
pacing
with
pedometer).
from map measurements.
5.6
day ranges
to my
calibrated
gave similar
in
0.5 km
a measured
distances
methods
accurately
estimated
10 full
for
a pedometer
between
15; p<0.01),
were
out
straight-line
The three
The correlation
n=
distances
by the use of
on the map.
be measured
could
calculated
by summing the
= 0.994;
map (and
distances
following
secondly
and thirdly
of distance
an
throughout
These plots
photographs,
were
paces while
locations
on
the baboons moved in a straight
that
aerial
they
known troop
sometimes,
chapter.
day ranges,
length,
to
an hour, and occasionally
In these cases the track
establish
Firstly
ways.
the
map. -Occasionally
or,
and relatively
were
known locations,
between
maps made from
and then
stride
to
order
hour
half
2) whenever
to within
no relief
troop
between known points.
in this
or
a tree
more than half
velocity
low
and
troop
by climbing
was not known, and I assumed that
the analysis
of
the
necessary
my position.
constant
to locate
bearings
on the
for
out of sight
ground
compass
the
recorded
the
on
intersecting
by interpolation
estimated
features
geographical
area helped
if
of
range
visible
position.
landmarks,
visible
of the
field
true
homogeneous
The
the
on maps (Chapter
was recorded
(ecotones)
borders
habitat
less
troop
the
of
between
measures
and map measurement
4000 paces
equivalent
The distances
for
to
all
5.4.2: Defining
Home Range
(1943)
defined
home
individual
in
normal
activities
for
but excluded
Burt
young",
its
go and occasional
(1975)
by Brown
earlier
which
that
used by the
area
is
alternative
use,
long-term,
intensive
to
define
for
to
define
home range
some
a primate
of
reviews
of the methods
the
et
I defined
by a taut
Sigg
and Stolba
their
troops
troops
into
every
definition
were
by the
said
quadrat
(in
within
the
to
have been
was tallied.
(1966)
the
5.7
1975,
or
in
to estimate
order
superimposed.
baboons
with
day's
were
a large
never
sample
the
home range
into
in order
to examine
the use of
of
quadrats
entered
Tinline
and
of each full
track
the
which
the
Hayne (1949),
such areas
and all
into
Lacunae
of home range.
of
influences
and Voigt
Firstly,
borders
all
very
course
Waser
including
home range,
Secondly,
baboons,
in
no
many comprehensive
are
have shown that
press)
of
are hard
the
further
There
case.
string
areas
is
have tended
during
1972,
second
or
field-workers
entered
exploited,
known to go.
were not
home range
the
encloses
were no areas
activities"
two ways.
as if
followed,
the
in
was enclosed
method
"routine
is defined
and Stumpf
probably
this
species
as
1976).
area entered"
home range
it
short-lived
over
or
this
concerned,
Clutton-Brock
any given
While
had
area
activities"
life.
animal
area
1976, Harding
troops
there
the
monthly;
Mohr
its
practice
used to estimate
Although
size
of
the
area which
follow
terms
in
(1980),
al.
and in
"the
"home range"
meaning
very
group,
(eg
et al.
The way in which
(1980).
in
the
Furthermore,
studies.
Hamilton
MacDonald
for
perhaps,
time
for
of
(1966)
as the
either
routine
might
was reiterated
Jewell
although
its
of
course
realistic
definition
the
and caring
mating
or young animals
be defined
pursuit
in the
animal
except,
specified
seasonal;
"in
presumably
practical
(1970),
home range
travelled
an animal
migrating
This
and Orians
that
recommended
which
by
traversed
area
food-gathering,
individuals.
of
and Brown
"that
as
of
into
areas
sallies
range
by the
transected
by the
baboons
were therefore
which
track
of
and each entry
excluded
from this
5. 'I. 3: Measuring Home Range Area
The accuracy
the number of
of the
with
full
area
and the
are
travel
several
Gi1Gi1,
Demment in
data.
range
against
This
towards
based
to
rise
to
the
first,
at
is
shape of curve
1978, and
Post
from
be estimated
new home
cumulative
been maintained.
has
after
that
animals
Amboseli,
observation
of
estimates
with
the
plot
on
the length
Normally
must usually
area
is
which
are
depends
was followed,
especially
(exceptions
steeply
This
an asymptote.
troop
samples,
method
for
can be estimated
home range.
Home range
time
the
the
of
a day
The normal
area
which
on small
prep).
tends
curve
size
kilometres
incomplete
area
for
day ranges
day ranges,
home range
home range
which
levels
it
which
off
by the equation
given
y=kx(1_e-mt)
The value
first
k of
just
of
data.
would
new areas
to enclose
whether
determine
is
the
are
all
the
animals
of
is
the method
the ground
almost
t to find
invaded.
If
are
or not
we believe
normally
construct
day
fits
the
Metzgar
and
to the
A, B, C or
sequence
in
C are
A, B and
ranges,
any common ground,
travel
but A happens
two, then
the chance
B,C,A or
C,A, B will
we have reached
that
of
method
by the other
enclosed
seen
the
at
on the
area
of m that
the value
thus
area;
home range
sensitive
three
enclose
to
home range
predict
rationale
B and C barely
whether
to
day ranges
Unfortunately
such that
arranged
to be the
taken
seem possible
essence,
(1974).
the
which
in
is
asymptote
enough full
This,
Sheldon
of
it
glance
basis
the
the
to
asymptote
the curve.
Researchers
this
sequence arrives
between the area predicted
the asymptote
asymptote
The
method
to
establish
confidence
in
the
order
at a more accurate
sequence would. ' The true
arbitrary
plot
(eg Altmann and Altmann 1970), but there
data were collected
to suppose that
the
is as low as possible
is no reason
than any
lie
somewhere
that
the day ranges in such an order
and that
order
the
which
result
home range area might
by taking
in
which gives
as high an
as possible.
method
adopted
derive
a straight
confidence
limits
here
limits
were then
was
line
for
to
use
of
known
both
treated
5.19
Metzgar
slope
parameters
as if
they
and
Sheldon's
(1974)
and
intercept
and
(Pollard
determined
1977).
the smallesE
to
These
ar'A
largest
home range
predicted
The most
from
Gilmer
et
al.
1975,1977,
1954
MacDonald
of
proportion
70% confidence
the
(1980)
ellipses
to the home range
in
the
records
the
constant
crossed
moved between
speed.
The
It
In order
habitat
type
track
which
the position
Principal
which
of that
the
the
habitats
baboons first
were
the day.
If
also
the baboons had left
riverine
trees
they had been out of the sleeping
enter
riverine
later
gallery
and humidity
of the field
the baboons.
with
for
of
habitat
leaving
each
habitats,
the
quadrat,
the
the
the first
forest
from
trees.
the sleeping
median
and the probability
half
of time
was calculated
determine
gallery
from any
and exit
by
time
that
the
fifth
of
by the time that
to
they
hour
said
were
an
time when they moved back into
in the day.
Temperature
centre
forest
for
at
baboons
the
which
by the end of the first
habitat
that
known track
the
The percentage
was recorded.
used
by the
by assuming
point
various
area
a habitat,
entered
in
spent
to
Trevor-
recorded.
greatest
quadrat,
a given
the animals.
movement
time
of
were similar
ch; Pmvnks
of
with
every hour after
baboons would have entered
it
of
area
points
was also
the
covered
of the troops
surface
enclosed
along
which the baboons were in each principal
for
1969).
to time the baboon's entry
to estimate
habitat"
"principal
speed
assumed
was seldom possible
habitat.
half-hour
for
account
each half-hour
for
technique
the
(Mazurkewicz
by radio-tracking
baboons
in their
which
home ranges
the
around
area assessed
each quadrat
the
showed that
Speed of movement was calculated
that
capture,
of
points
variance
and Hackett
Deutsch
and
been
has
et al.
1972, Stickel
1969, Metzgar
1980)
1943,
1975, Burt
1968, Koeppl
1965, Jorgensen
Hackett
so far
emerged
(Brown
animals
1980, Mazurkewicz
al.
et
around
the
of
and Barbour
1973, Harvey
ellipses,
trapping
repeated
Trevor-Deutsch
and
drawing
the
have
to
method
sophisticated
statistically
of
studies
and Home Range Area
Ellipses
5.4.4: Confidence
area.
were recorded
area by other
air
temperature,
5.9
at
members of SAPP while
Speeds of movement could
the prevailing
hourly
therefore
the
camp in
the
I was following
be compared hour by hour
sun temperature,
humidity
and vapour
. _..
0
5.1:
Figure
in known home range
Increase
of Anecdote's
sample size.
kilometers)
(400 quadrats
- 16 square
troop
with
increasing
40
. r4
cd C)
Lý t,A
'C C
cet co
201
00
Q.
0
zý4
Sample
Size
(number
of
full
day ranges)
`
,.
r
5.2:
Figure
in known home range of Fable's
Increase
sample size.
(800 quadrats
kilometers)
a 32 square
troop
G)
bD
O
E
O
800
O
N
400
it
'O
ai
4- .
O
G)
A
E
z
Sample
Size
(number
of
full
day
ranges)
with
increasing
deficit.
pressure
methods used in the examination
Some of the more detailed
behaviour
in the appropriate
are given
of ranging
in the results.
sections
RESULTS
of Home Range
5.5: Size
the
troop
Anecdote's
For
of
plot
taken
the number of day ranges,
home range against
in the known
the number of quadrats
fits
chronologically,
the curve
e-0.11t)
Area = 428 x(1where
home range
estimated
this
(16.7
area
For
results.
similar
in
quadrats
(Fig
line
straight
allows
Anecdote's
Fable's
troop,
home range
= 24;
18.8 sq km (Fig
or
full
day ranges
both
plot
of
number
against
an
closely
numbers
of
day
on
covered
gave
methods
the
however,
5.1).
to be set
limits
group
an
ns) giving
confidence
Superimposed
for
known
the
470 quadrats,
method
sq km).
24.2
to
sq km Thus
18.8
of
area
(1974)
= 18.1; df
squared
about
of
area
and Sheldon's
Metzgar
(chi
number of day ranges
t=
+ 41
of
is
ranges
a
5.2)
Area = 28.On + 45
(r = 0.96; n= 26; p <0.01).
Increasing
tendency
method
these
quadrat
the
of
line
shows that
data
up
an asymptote.
smallest
home range
52.8 km sq, compared
A final
day ranges.
the cumulative
and Sheldon's
Metzgar
area
prediction
new area enclosed
which,
together
with
small
the 42.7 sq km given
by
km, while
Fable's
on the
of home range area was
within
full
day ranges.
the fourth-last
of 22% in known home range area was made,
increases
curve to tend towards an asymptote
In summary, Anecdote's
(1974)
of
The data tend towards an asymptote of about 40 sq km until
day range, in which an increase
the
basis
possible
with
to
difference
km made little
reach
be about
full
made by plotting
to
the
would
superimposing
size
1 sq
to
in
the next
three
days caused the
68
at about
sq km,,
known home range covered an area of about 20 sq
covered at least
42 and probably
5.10
45
km
than
sq
more
Figure 5.3(a):
Known home ranges of two troops
of P. papio at Mt. Assirik.
(roughly
troops
9 sq km) is shaded.
both
by
Zone shared
(shown
follows:
black
dots)
as
are
Sleeping
numbered
as
sites
I Xanadu
2 Elephant Rib
3 Secret Valley
4 Assirik
Valley
5 Flycatcher
Falls
6 Old Camp
(camp)
7 Amphitheatre
8 Cross Valley
0
9 Stella's
Waterfall
Figure 5.3(b)s
troop with sleeping
Known home range of Fable's
sites
is
"Capture"
home
Area-of
range
confidence
ellipses.
by the 90% confidence
ellipse.
closely
approximated
Known
e1 F0
Wilk S
Copy
and
most
The home range area of Fable's
but by the 90% confidence
Anecdote's
and drainage
streams
Assirik,
where
Fable's
troop's
containing
its
its
lying
areas
range,
containing
Anecdote's
troop
to
cliffs
did
the
and
ravines.
between
not shift
Anecdote's
troop,
species
its
with
largest
had the tenth
(Kendall's
ranges
5.6:
(Fable's
baboons
troop,
0.13,
ns;
day by Fable's
range
of
which
fragmentary
for
than all
(Table
I have data
5.1).
home range of 18.8 sq km,
including
known troopsq
troops
larger
but 3 of the
Fable's.
home
larger
to exploit
followed
were
14 in
the
wet
for
total
a
9 in
season,
The distribution
3.03,
kurtosis
troop
dry;
and by Anecdote's
ranges
14
troop,
lengths
The mean distance
km,
day
full
Anecdote's
day range
of
ns).
was 8.1±2.1
the
of
49
was normal
in
travelled
7.6±2.1
troop
a
km
5.2).
In neither
significant
1.3
more
tau; p<0.097).
and 12 respectively).
(Table
included
home
dry
the
Movements of Troops
The
(skew
in
the
Studies
of baboon for
trend
known
was larger
home range of all
There is a non-significant
The
into
while
Mt. Assirik,
bases.
their
season
Mt. Assirik,
of
other
troop
possibly
wet
quadrats
of
at
season
the seasons.
with
The home range area of Fable's
of other
of
environs
proportion
the
common
dry
the
of vegetation
in
west
in
this,
high
a
the
5.5.2: Home Range Area: Comparison
30 troops
Despite
were
and
to the west of Mt.
largely
ranged
range
to
confined
quadrats
troop
dense thickets
extended
Ridges
and marigots.
included
their
on Mt. Assirik,
was centred
relief.
range
with
undifferentiated
season
Fable's
was little,
home
cliffs,
plateaux
rare.
there
troop
Fable's
known home range
many laterite
by the 70%
was not approximated
(Fig 5.3).
ellipse
troop's
included
hence
troop
respectively).
difference
troop
between seasons,
difference
respectively)
nor Anecdote's
Fable's
or
Similarly,
between troops
distance
mean
there
within
was
seasons
5.11
was there
in either
travelled
no
(wet
a statistically
variance
(t
(F = 2.0 and
= 0.8
statistically
F=1.5;
season
and
significant
t=0.8;
0.4
Table 5.1:
Home Range Areas of Troops of Baboons
Square roots
Species
,
log-normal
home
range
areas
are
of
Area
Source
0.9
3.0
Dunbar and Dunbar 197lc
Ransom 1971
3.2
Suzuki
P. anubis
1979
3.9
4.3
5.2
6.3
6.0
19.7
24.8
Rowell 1966a
Aldrich-Blake
et al. 1971
Rowell 1966a
Maxim and Buettner-Janusch
Nagel 1973
Harding 1976
DeVore 1965a
52.0
73.0
Popp 1978
1959
Boiwig
16.7
40.0
43.0
61.0
Altmann and Altmann
Post 1981
Rasmussen 1979
Oliver pers comm
1963
P. cynocephalus
1970
P. ursinus
Hamilton
2.1
zI. 0
4.7
4.8
197691978
et al.
It
6.5
9.1
9.4
10.7
12.7
Hall 1963a
Hamilton et al.
Hall 1962a
if
13.0
14.8
1970
Saayman
Stoltz
and
1963a
Hall
1978
33.7
P. hamadryas
15.0
29.0
Nagel 1973
Sigg and Stolba
18.8
42.7
This
This
P. papio
dry
season
ranges
Assirik
F=1.7;
included
travelled
t=0.3).
(F = 1.0;
about
study
study
in press
(Anecdote's
troop)
(Fable's
troop)
and none between
t=0.9).
7.8±2.
Okm per
5.12
In
summary,
day
troops
with
the
baboons
irrespective
of
all
day
at
Mt.
season
or
Table 5.2:
in Two Seasons
by
Troops
Two
Ranges
Day
of
Lengths
Troop
Rank {
Anecdote
Fable
Wet
Dry
Wet
Dry
{ 4.6
5.5
{ 6.8
3.9
5.0
7.2
5.3
5.4
5.5
3.5
5.2
6.1
4
{ 7.2
7.3
5.7
6.4
5
6
7
8
9
10
11
1 8.3
{ 8.5
1 8.5
18.5
{ 9.5
{ 9.6
19.7
7.8
8.1
8.5
10.8
11.0
6.2
6.6
7.5
7.8
8.2
8.8
9.1
7.2
7.4
7.4
7.4
9.4
9.4
9.6
12
110.1
9.5
9.8
13
14
{10.1
110.5
9.9
13.1
x
{I 8.38
7.68
7.75
7.40
sd
1 1.79
2.51
2.21
1.95
{
1
2
3
{
troop.
The distance
from
predicted
if
general,
by time t (t
the
(0.347
the
between
asymptotically
one standard
ranges
4.3±1.8
Anderson
Stoltz
than
km;
travelled
at
(1 -e-0.3
of
baboons
Dunbar
et
day,
for
n:
at
had
reported
5.13
to
tend
Mt. Assirik
aj..
where
r
is
the
15,
x (t-9))
apparently
and Dunbar
(1970)
Saayman
and
that
so
and P. cynocephalus
Aldrich-Blake
1980b,
day
travelled
distance
final
of the mean day range
deviation
the
time
xd+0.3
and the
of
Mt. Assirik
P. anubis
did
time
and t the
coefficient
genus.
In
midday.
after
distance
the
d
range and
prediction
with
The population
the
at
x (t - 12) + 0.089)
r=
within
any time
be
could
ranging
> 12: 00) then
The correlation
correlation
day's
a day's
end of
travelled
distance
1 was the predicted
1=
improved
by the
travelled
(around
have
(mean of
1971,
Altmann
1974o,
Harding
that
a mean day
troops
and
of
1976,
range
6.5 km) for
day
shorter
5 estimates
Altmann
1970,
Nagel
1971).
one population
of
P. ursinus
ranged
for
estimates
1979,
P. ursinus
are
and Stolba
in
troops
of other
km per
(Hall
1962a,
1963a).
further
than P. papio,
Mandrillus
1975).
sphinx
Of all
1973,
a
Stolba
ranges
length
for
10 day ranges
with
the
with
also
from 0.6 km (T. gelada,
(P.
1974c)
11.1
km
hamadryas.
Dunbar
to
and
other
although
Nagel
Mean day range
of baboons varies
species
day,
km (Kummer 1968a,
press).
(Jouventin
8
km
day
each
about
8.0
ranges
9.1±2.1
being
estimates
Sigg
lower
P. hamadryas typically
baboons only
mean of
mean of
a
over
over
single
Dunbar
included,
Kummer 1968a).
5.6.1:
Speed
There was no significant
day
ranges
which
at
movement
correlation
was
for
examined
(rho)
coefficient
19 of
a sample of 24 full
between
autocorrelation
(Table
intervals
half-hour
successive
in
correlation
speeds
The
5.3).
of
common
was about 0.3.
Table 5.3:
Autocorrelation
Sample
Fable's
Troop
Number
r
Sig
level
r
489
.
5%
680
637
1%
1
197
.
413
.
2
3
4
. 355
5
Anecdote's
.
5%
320
298
.
421
.
164
.
1B7
-.
179
.
093
.
298
.
10
11
12
1%
307
.
241
.
036
.
303
.
9
Big
level
209
.
.
187
.
455
.
214
.
Troop
341
.
342
.
.
6
7
8
Intervals
of Speed at Half-Hour
5%
Over the whole year the median speed of movement of a moving troop
recorded
0-2.8
on the
kph)
for
half-hour
and for
There was no difference
was,
Anecdote's
in
the
for
troop,
Fable's
0.66
troop,
kph
median speeds with
5.14
0.72
(range
which
kph (range
0-2.8
kph).
the two troops
(medians
moved
the
season
dry
the
df
chi
median
the
season
median
(1970)
Altmann
kph)
0-3.9
speeds
kph
0-4.0
(median
wet
in
kph;
7.48;
=
squared
0.67
of
a median
speed
season
(medians
test,
by Altmann
reported
kph,
0.44
about
of
max speed
over
and
range
(mean of
P. hamadryas
for
recorded
1979;
Stolba
these
than
higher
those
to
kph,
was 0.80
chi
dry
the
the
ns).
are
similar
troop
at
In
ns).
test,
moved
in
P. cynocephalus
for
but
range
1;
=
df
= 0.072;
These
troop
0.65
and
(medians
kph
Anecdote's
wet
squared
0.64
= 1;
df
Fable's
of
movement
of
was
p<0.01).
in
kph,
it
= 2.51;
squared
chi
speed
season
= 1;
kph
test,
4.7
hour
one
0.8
kph,
Kummer 1968a).
Speeds
recorded
from
estimated
sightings
The
variable.
was
quadrat
Anecdote's
the
3.9
across
was seen
quadrats
respectively.
high
the
(Note
speeds
troop
5.6.2:
that
since
is
the
the
a given
time
lay
quadrats
3.1
quadrats
across
interval
about
biassed
are
more quadrats
while
kph
2.9
and
between
are
a
in
were seen to
troops
were
season
across
speeds
which
3.6
about
at
the
across
season,
wet
a quadrat
with
dry
median
fast.
moving
fastest
distributed
1.1
kph
and
towards
crossed
if
were
not
)
with
speeds
(chi
in the middle
the
first
third
periods
three
of
for
the
day
(chi
which
the
baboons were
fell
more frequently
than
(chi
squared = 7.57;
df = 2;
chi
to occur
Second fastest
p<0.01).
5.15
in
into
the
4.08;
squared =
third
last
df = 2;
stationary
the
00-15: 590
central
in
p<0.025).
half-hour
on the
third
but
to fall
speeds tended
squared = 8.92;
expected
moved
6: 00-11: 59,12:
goodness-of-fit,
squared
of the day.
baboons
periods
Top speeds tended
p<0.05).
not
the
which
the
randomly across
16: 00-19: 59
df = 2;
while
in
speeds
speeds
in
more
Time
The
and
in
troop
were
troop
30
often
were
hence
and
either
for
mean speed
quadrats
of
movement
move across
The fastest
Median
kph.
to
across
intervals,
shorter
Fable's
by
the
represented
movement
recorded
kph,
troop
hour
of
at
fastest
same season.
move
1.3
Speeds
travelling.
minutes
half
on the
of
the
day
Riverine
on laterite
the
tended
(Table
trees
tended
to
habitat.
in
occurred
percentage
of
the
forest
these
first
were
grassland
Table
and
to
fifth
first
first
of
habitats
entries
to
made in
the
were rarely
early
open
first
day
to
corresponded
forest
in
the
woodland
fifth
of
or never entered
the
reflected
the
than
day
(Table
and
2/5
the
day,
in this
of
5.5).
these
of
baboons
the
half
to
entries
while
which
Roughly
the
of
other
any
skewness
probability
left
baboons
habitat
given
a
the
areas
earlier
to
visits
the
of
social
scrub
baboons
the
after
the
dry
Combretum
associated
shortly
had left
and
woodland
The
closely
entered
Once they
5.5).
open
the
and the
be first
to
enter
distributions,
getting
forest
gallery
dry
and scrub
scrub
period.
5. ':
Visit
Median Time of First
Whole Year; Both Groups
Day Ranges
Data From All Full
Habitat
Median
Time
Riverine Gallery Forest
Combretum scrub on laterite
07: 00
08: 15
Open Woodland
09: 30
Dry Forest
09: 30
Scrub
10: 30
Grassland
11: 30
12: 30
Scrub
Grass on laterite
Table
5.5:
Percentage of Observations
on Which First Visit
1/5th of Day
To a Given Habitat was in First
Habitat
Troop
Fable Anecdote
Season
Wet
Dry
Total
86
97
92
80
86
Combretum scrub on laterite
Open woodland
Dry Forest
57
32
56
71
64
40
58
39
32
71
59
50
64
48
39
Grass on laterite
Scrub
Scrub Grassland
20
0
0
11
19
0
16
6
0
16
11
0
16
9
0
Riverine
Gallery
Forest
5.16
5.6.3:
Climate
There
was
by
travelled
troops
the
within
either
no
the
and
or
seasons
between
the
temperature
of
relationship
significant
maximum air
the
across
(r
year
distance
> 0.3;
never
the
day,
p
never
< 0.05).
There
were
either
humidity,
or
vapour
5.6.4:
from
590 positions
of
the
troop
of
the
troops
The probability
p of
site
was
inversely
and was given
by the
sleeping
site,
The median distance
they
chi
to
troops
correlated
in
the
frequency
sleeping
Anecdote:
distance
«
sleeping
with
22%; Anecdote:
Both
by
movement
temperature,
in
coefficients
no
slope
but
the
in
of
visited
the
the
troops
to
quadrat
inverse
more
(medians
frequency
was
quadrats
from
the
used any quadrat
the
which
linearly
also
nearest
sleeping
variance
(Fable:
test,
with
square of the distance
much of
than
sites
sleeping
evenly
The
p<0.001).
seasonal
1.9 km. The baboons
was about
home ranges
their
of
either
closer
was 0.78 km,
site
to the
in
the
wet 21%, dry
wet 20%, dry 21%).
troops
sites
their
explained
site
site
significantly
seasons
0.001),
which
from a sleeping
any point
sleeping
distance
that
from
distance
their
a
xd-0.02
df = 1;
both
to
d from
distance
a given
at
equation
from
used
was recorded
site
day ranges.
troop
of the troop
they
the
with
(p always
nearest
sun
sleeping
full
during
linear
be found
squared = 54.88;
nearest
proportional
nearest
be if
would
both
site
the
to
were normally
temperature,
(correlation
the
to
finding
the median distance
home ranges
of
zero).
p=0.06
while
speed
Sites
The distance
for
pressure
hourly
air
deficit
different
Sleeping
with
season
either
significantly
case
correlations
no
in
troop
between
tended
to
move faster
in the wet season (Fable:
0.1
from the site
kph/km,
p<0.006),
when they
slope
but
0.17 kph/km;
there
on speed in the dry season.
5.17
were
further
from
p«0.001;
was no effect
of
5.6.5:
Habitat
The estimation
of
of movement across
on
the
and frequency
proportion
troops
(Table
p<0.01)
habitat
quadrats
(regression
2.
Chapter
in
present
the
the
by each habitat.
affected
the speed with
(two
28 cases
lines,
which the
two
troops,
660
between
n
quadrat.
speed of movement
covered
of
Speed
were regressed
entirely
13 out
in
a quadrat
habitats)
seven
seasons,
into
habitats
a quadrat
of a given
moved across
in
of entry
various
into
of entry
The percentage
described
were then used to predict
lines
These regression
and frequency
the
of
is
type
habitat
2250,
and
5.6).
5.6:
Table
Line
from Regression
Speed (kph) Predicted
100% of Habitat
Across Quadrat Containing
*p<0.01
*-p<0.001
Troop Season {Bush
I12.05*
FW
D
12.22**
11.59
D
10.98
I
AW
Scrub
Habitat
Woods
Combr
1.74**
1.610
1.51
1.26
1.50
1.28
0.87
1.33
Forest
Grass
Gallery
1.18
0.94**
1.34
1.08*
1.540
1.20
0.990
1.30
0.7700
[-0.3]**
[-0.1]**
[-0.2]**
1.15
1.00
1.04
[-0.3]00
[-0.2]
11.71
1.03
1.21
1.43
1.15
1.34
f
KEY: F= Fable's troop
A= Anecdote's troop
D= Dry season
W= Wet season
Scrub = Scrub
Bush = Scrub grassland
Mean Speed
Woods = Open woodland
Grass = Grass on laterite
Combr = Combretum scrub on laterite
Forest
= Dry forest
Gallery
Gallery
Forest
= Riverine
In both seasons both troops
gallery
forest
the
more scrub
dry
season
quadrat,
slower
plateau
in
movement of
and scrub
grassland
the
moved faster
but
the
was present
in the
it
in
the
troop
a
wet season,
moved.
quadrat
both
troops
quadrat.
there
the
was negatively
the
5.18
wet
troop
Fable's
was in
more dry
Increasing
in
the more riverine
tended to move slower
the
forest
more dry
correlated
season.
quadrat.
was present
forest
proportion
moved faster
In the
in
was present,
with
This
the
on laterite
grass
of
the
the
trend
speed
of
was also
though
present,
not
Scrub on laterite
statistically
was never statistically
It
the
(Table
5.6).
scrub
grassland,
The troops
have
and I
quadrat
Almost
(two-way
anovar
forest)
but
speeds
with
multiple
in
speeds
of
Riverine
movement
1.0
between
accounted
troops
in
both
little
for
3
multiple
gallery
through
higher
on
laterite
of
the
by
than
the
(Scheffe's
in
and
speed
the
habitat.
single
any
in
scrub
variance
6.4%,
and
forest
riverine
moved
more variance
seasons
("Bush")
more frequently
more dry
the
In
present.
frequently
the
wet
with
scrub,
than
did
any
a single
habitat
scrub,
which
enter
a quadrat
The
is
taken
variance
quadrat
to
the
accounted
quadrats
with
mean that
entirely
frequency
by
5.19
in
the
scrub
and
forest
were
quadrats
more
in the quadrat.
completely
a frequency
p<0.001)(Table
frequency
for
entered
was present
that
predicted
anovar,
consisting
in
troops
on laterite
a negative
predicted
both
in the quadrat,
gallery
and riverine
would be entered
(two-way
on the habitat
forest
season
lines
quadrats
entered
the
frequently
less
and scrub were present
the more grassland
The regression
given
grass
between
for
with
troops
accounted
with
more scrub-grassland
lines
a
habitats.
Both
with
in
slowly
gallery
significantly
Very
be
could
the
was
p<0.05).
and 5.0%,
forest,
Scheffe's
p<0.025;
through
moved
forest,
gallery
Tables)
the
comparisons,
pure
The
gallery
moved very
p<0.025
at
which
with
they
which
19.7;
=
significant
speeds
("Bush"
grassland
other
the
of
forest.
were due to riverine
effects
habitats
habitats
all
habitat
each
a quadrat
riverine
have
would
in
gallery
in
speed
of
habitat.
this
F between
quadrat
riverine
baboons
the habitat
all
comparisons,
the
only
containing
in
a negative
that
a pure
the speed that
line
have moved fastest
slowest
predicted
assumed
in
have
to
would
and
line
regression
which the regression
from the regression
to predict
be expected
might
for
season.
significant.
was possible
troop
habitat
was the only
plateau
the dry
in
so,
significantly
5.7).
covered
which depended
The regression
(Bush)
grassland
baboons would
and in
be unlikely
to
habitats.
these
one
of
of
with
which
habitat
was
the
never
baboons
more
entered
than
any
14.5%.
Table 5.7:
Frequency
of Entry into Quadrats
Predicted
Frequency
of Entry into Quadrat
100% of Given Habitat
When Quadrat Contains
p<0.01
p<0.001
*+ý
Season I
Troop
Scrub
Bush
1 0[-. 13]** 0[-. 03]**
10[-.
19]**
0. 02**
1 0[-. 37]**
0[-. 14]**
10[-.
0[-. 14]**
20]**
1
Numbers in square brackets
Fable
Wet
Dry
Wet
Dry
Anecdote
Different
habitats
(Friedman's
scrub
S;
accounted
with
Table
particularly
forest,
gallery
riverine
0.81**
0.23
0.46
0.35
0.68
0.54**
0.75
0.94*0
9.
6.
10.
9.
0.39**
0.16
0.64**
0.35
taken
are
df
to
imply
Combretum
in
variance
frequency
the
and especially
scrub,
while
more (Table
for
variance
of
p<0.005).
the
of
01**
62**
07**
88**
zero.
amounts
6;
=
a quadrat,
accounted
1.15**
0.42**
0.79**
0.51**
differing
little
baboons entered
the
which
Gallery
= 27.25;
squared
for
Grass
for
accounted
chi
Habitat
Combr Woods Forest
5.8).
5.8:
Frequency of Entry into
Percentage of Variance Explained
Seasonl
I Bush
Troop
Fable
Wet
Anecdote
5.6.6:
available,
Scrub
Grass
Gallery
1.0
3.9
0.6
0.4
3.7
0.1
10.4
i
0.8
1.1
0.0
0.1
0.9
0.6
14.5
Wet
Dry
I
0.9
2.4
2.3
0.0
0.0
0.3
0.0
0.7
0.4
0.1
1.0
7.9
13.0
Neither
(Table
Habitat
Combr Woods Forest
Dry
1 0.5
i
Time Spent
as those
Quadrats
by Given Habitat
troop
habitats
5.9).
in Various
Habitats
shared
their
out
occurred
The
forest,
and grass
by both
troops
field
time
index,
on laterite
in both
the
of
ratio
a preference
in
(Kendall's
area
spent
shows that
to
riverine
while
5.20
proportion
tau = 0.14 ns)
proportion
were used consistently
seasons,
in
time between habitats
gallery
of
habitat
forest,
dry
more than expected
Combretum on laterite,
scrub,
and
5k
(Bush) were used consistently
scrub grassland
35 times
as much time
spent
nearly
forest
and about 5 times
Table
less
less.
in
as expected
gallery
riverine
(Table
in scrub
than expected
the baboons
Indeed
5.10).
5.9:
in Various
Time Spent
Principal
Habitats
Percentage of records of troop
habitat
in quadrat with given principal
Troop
Season
Fable
Wet
Dry
Woods Gallery
Forest
Anecdote Wet
Dry
Grass
Scrub
Bush
39.0
1 30.0
22.1
28.9
11.3
17.5
13.1
7.9
7.0
11.4
5.6
4.4
1.9
0.0
134.9
125.4
I
25.3
31.2
16.1
10.7
11.8
8.8
5.9
17.6
3.8
4.9
2.2
1.5
32.5
26.7
13.5
10.6
10.5
4.7
1.5
20.8
29.5
24.9
3.6
Mean
%
3
Rank
% in field
KEY:
area
=
Forest
= Dry forest
Gallery
gallery
= Riverine
Grass = Grass on laterite
Table
Combr
0.4
14.9
5.8
1
7
4
5
percent
in field
habitat
area
of principal
Woods = Open Woodland
Combr = Combretum scrub on laterite
Bush = Scrub grassland
forest
6
2
5.10:
Ratio
of
Available
to Habitat
Time Spent
Forest
Woods Gallery
Combr
Grass
Scrub
Bush
1.88
0.75
28.25
0.88
1.21
0.22
0.53
1.44
43.75
1.97
0.18
0.00
0.86
40.25
0.53
1.68
0.98
0.79
1.02
0.15
0.61
1.22
1.06
26.75
0.59
3.03
0.20
0.42
Mean
1.55
0.91
34.75
0.39
3
1.81
2
0.19
Rank
0.70
5
Troop
Season
Fable
Wet
I
Dry
1
Anecdote Wet
{
Dry
4
1
6
7
KEY: see Table 5.9
5.6.7:
Topography
Topography
selection
ratio
as follows.
topography
was assessed
for
the
the
each topographic
The proportion
in
by
seasonal
method
type
of the entries
given
into
all
Chapter
2.
A
for
each troop
quadrats
of a given
was calculated
home range was divided
5.21
in
by the proportion
of
quadrats
less
than
troop
the
one
less
type
frequently
home range;
that
contained
which
than
entered
quadrats
expected
on the
chi
squared
in
fastest
(at
quadrats
1.4 kph)
about
(about
water
season
= 9;
0.75
was
topographic
in
frequency
by
marigots
in
and slowest
the
in
or
sites
941.5;
S=
baboons
the
quadrats
or by sleeping
kph)
(Friedman's
with
p<0.005),
characterised
quadrats
its
of
by topography
Speed of movement was affected
df
that
containing
basis
ratio
selection
more frequently.
when greater,
= 31.38;
When the
topography.
moving
undifferentiated
by dry
characterised
(about
0.6
kph)(Table
5.11).
Table 5.11:
Speeds of Movement of Two Troops in
Topographic
by Various
Characterised
I
Wet
Dry
Anecd
Wet
Dry
Undf
CaSt
Plat
Clif
Ridg
DrSt
Ravi
DrWa S1Si
1.18
1.68
1.33
1.51
1.33
1.21
0.96
1.45
1.11
1.03
0.96
1.07
1.26
1.33
0.89
1.07
1.03
0.73
0.52
0.44
1 1.47
; 1.20
1.54
1.00
1.26
1.16
1.26
1.21
1.19
1.10
1.19
1.10
0.84
0.68
0.77
0.82
0.56
0.63
0.70
0.47
1 1.38
1.35
1.24
1.22
1.11
1.08
1.03
0.89
0.74
0.59
df = 9; p<
0.005
I1
Mean
Friedman's
Feature
Topographic
Troop SeasonI Mgot
Fable
S = 941.5;
Chi squared = 31.38;
CaSt = Catchment Stream
Ridg = Ridge
Undf = Undifferentiated
Clif = Cliff
Key: Mgot = Marigot
Plat = Plateau
Ravi = Ravine
S1S1 = Sleeping
Stream
DrSt = Drainage
DrWa = Dry Season Water
Site
of topographic
For definitions
(Friedman's
The effect
S=
squared = 7.4;
74; chi
of topography
by the
vegetation
this
so is
is
2.
growing
suggested
in
by the
the
high
in which both habitats
in
order
of
the
df = 3;
ns).
on speed of movement was probably
of Table 5.12,
descending
see Chapter
types
depend
did
topography
upon season or
not
on speed
of
The effects
troop
Quadrats
Features
different
percentages
which
the
types.
That
near the main diagonal
and topographic
speed with
5.22
topographic
mediated
types
troops
are arranged
moved through
them.
Table 5.12:
Percent
by given Topographic
characterised
uadrats
ich was covered by a given Habitat
of all
in each topographic
Major habitat
Habitat
Combr Woods Forest
Scrub
Bush
Type
Topographic
is underlined
type
Gallery
Grass
Marigot
0.0
2.5
38.2
19.6
23.0
16.5
0.2
Undifferentiated
Catchment Stream
Plateau
9.9
30.3
12.6
27.3
16.9
3.0
0.0
12.8
0.2
30.1
1.1
3.9
48.1
38.7
8.9
12.4
15.5
2.2
26.1
0.0
0.1
3.4
9.3
10.4
20.0
16.6
3.0
33.3
44.1
28.0
22.8
8.2
0.8
0.2
0.0
8.5
27.6
5.1
23.2
32.2
3.2
0.2
0.0
4.0
0.5
0.4
13.8
1.2
19.4
15.4
18.1
11.2
10.2
4.9
39.6
35.4
28.2
20.8
15.0
13.9
8.6
6.2
33.2
each
troop
Cliff
Ridge
Drainage
Stream
Ravine
Dry Season Water
Sleeping Site
The
frequency
on the
depended
seasons
never > 0.025).
those
for
sleeping
sites
type
Shade and Visual
Shade and visual
be
similar
of that
habitat
regressed
manner.
on the
in
in
ravines
p
with
for
any
multiple
5.13).
in
Combretum scrub
habitat.
percentage
each
each habitat,
the quadrat
Speed
in
movement
of
product
and the mean proportion
Mean visual
of
in
quadrat
the
and
density
frequency
shade and degree
quadrats.
5.23
were assumed
on
grass
and
of
each
of
2.
Chapter
in
grassland
and scrub
scrub
mean shade
that
(Scheffe's
was measured as described
density
by summing, for
dense shadow in
similar
for
those
than
higher
squared,
and 3.61,
between 0.27
both
Density
those
The
respectively.
calculated
to
(chi
quadrat
significantly
except
density
The shade and visual
to
being
of
quadrats
entered
that
varied
ratios
p never > 0.025)(Table
comparisons,
5.6.8:
topography
Selection
topographic
other
which
with
in
the
laterite
season
was
percentage
in
the
ground
of
was assessed
of
visual
entry
density
in
a
were
in
Table 5.13:
Freq uency of Entry into Quadrats Characterised
Various topographic
Features
Topographic
Anecdote
Fable
I
I
by
I
I
I
i
I
I
Plateau
0 sr I sr
II sr If
0 sr If
If
# sr If
I
I
I
I
I
1137 70 1.05 1 45 35 0.76 1118 48 0.94 188 195 0.27 10.76
Wet
Dry
1 67
1 48
1313 219 0.77
1 65 44 0.80
Marigot
117
Drain Str.
Ridges
Cliffs
Dry S. Wat.
Ravine
1235 125 1.01 169
1 67 41 0.88 1 26
1221 107 1.11 1113
I 54 16 1.82 1 38
140
12 1.79 141
1 57
15 2.90
I
KEY: f=
#=
sr
1 72
1147
1225
1 23
1 39
0.95
0.77
0.92
1.88
1.51
frequency
quadrats
of given
with
16 1.43 10.92
1.02 1 29
0.86 1141
1.14 1122
0.87 136
1.35 1 52
(min
11.00
11.06
11.15
11.68
11.94
13 3.27
13.05
1
1
was seen to
enter
topography
in
topography
from
home range
seasonal
/ (0/sum of 0's)
of f's)
in dense shadow at
of the ground in any quadrat
The mean proportion
1.02
1.73
1.43
2.16
3.12
17
49
51
10
10
171
10 3.61
troop
which
quadrats
of given
Ratio.
Calculated
noon was 20.0±15.9%
24 0.89 1 38
1
Observed
(f/sum
26 1.11 10.89
36 1.35 10.92
27
65
75
10
11
1 95
14 2.41
I
Number of
Selection
=
43
20
73
12
16
Dry
1 48
1 81
48 0.87
66 0.72
3 0.59 1 56
12 0.76 13
1 81
Site
Sleep
Wet
1110
1125
49 0.81
35 0.81
Undiff.
Catch Str.
I
IMean
Feature
1.7%, max 70.2%)
in
but only
the wet season,
9.3±9.9% (min 1.4%, max 53.9%) in the dry season.
In
the
the more shade they
frequently
dry
season
troop
apparent
the
variance
entered
troop
on
for
Both
20% of
for
which
the
the
in
with
which
had
quadrats,
(p < 0.05,
p<0.0001,
tended
to
enter
areas
in the wet season (p < 0.0001 for
Anecdote's),
across
in both
with
shade
with
was
no
while
2% of the
11% of the variance
in the dry season).
troops
frequently
troop,
Fable's
in both seasons
in the wet;
(p < 0.0001
frequency
the
movement
of
and in the
quadrats
entered
the quadrat
variance
For
speed
(p < 0.01),
troops
both
more
quadrats
entered
contained
shade in
quadrats).
moved slower
accounted
accounted
with
amount of
for
effect
Anecdote's
season,
frequency
accounting
Fable's
l
the
with
correlated
cases,
troop
Anecdote's
season,
wet
but visual
contrary
to
density
of
Fable's
had no apparent
expectation.
5.24
In
the
dry
visibility
poor
effect
troop,
p<0.01
less
for
on speed in that
season,
however,
both
troops
entered
areas
in frequency
variance
(p < 0.0001)
(about
density
visual
(p < 0.00001)
more frequently
10% of
and moved through
for)
accounted
the
(about
variance
in
differed
greatly
13% of
the
them more slowly
if
for)
speed accounted
the
was higher.
DISCUSSION
5.7:
Home Range Area
two troops
The
Anecdote's
area.
which
troop
troop
the
of
was only
to
well
ranged
the
its
"expand"
troop
It
troop.
the
is
north,
of
18 or
south
the
limit
and east
of
Stella's
apparent
might
sites
and the
known home range
of
and that
it
true
of
had
sites
station
research
its
while
The
sleeping
the
Thus
observer,
range.
three
that
possible
westernmost
the
home
of
the
many of
study.
known sleeping
known
or
the
two
of
one
8 sites,
in
new to
of
positions
relative
late
number
two
to
from
6 of
from
troop
areas
restricted
Anecdote's
been due to
home range
its
to
of
large
invaded
with
opportunity
have
the
known to be used by that
troop
restriction
was followed
home range
predicted
followed
always
almost
troop
continually
other
less
was
Fable's
were only
this
troop
sites.
sleeping
the
apparently
in
range,
Waterfall
sleeping
site.
The value
troop
during
Fable's
troop
covered
in any single
the
unlikely
one
year,
complete
that
but
be an underestimate,
may well
than
the home range area of Anecdote's
19 sq km for
all
and
the
study
year
of
home range
the less
secondly
of
year
The area
life
of a troop
1s probably
smaller
of
troop
5.25
home range
known.
the
existence.
the
of
was probably
the
commonly visited
the
most
troop,
parts
may shift
firstly
will
its
because
be visited
range
during
it
is
in any
its
5.8: Movements of Troops
Baboons
recorded
most
baboon
savanna
P. hamadryas.
day
Long
consequence
far
as
dry
troops
of
a
long
day ranges
season
and large
the
the
did
presumably
are
At Mt. Assirik
in
did
as
than
day
each
P. hamadryas
of
to to reject
length
range
they
in
in large
in smaller
wet season
compensating
for
than
troops.
of the two troops
it
the
linearly
did
in
in the
they
dry
season.
to the size of the troop,
kph, and a troop
(Fable's
figure
seasonal
Assirik
If
we find
is close
of
the
in
syndactylus
since
It
seasonal
seems possible
the baboons
were able
troops
remained
may have been
in the dry season.
faster
speed of
then interpolating
3) with
in the wet season
movement was related
between a dry season
a median speed of about
that
(Anecdote's
135
of about
members
0.65
to the observed
may have been related
forest
median time of "entry"
speed of 0.67 kph, and suggests
to seasonal
differences
the sleeping
sometimes
into
wet
should have a median speed of about 0.69 kph.
it
in the median speed of movement of the troops
difference
gallery
in day
have had to had they
size
the
to a troop
When the baboons had left
riverine
season
dry
of about 250 members and a median speed of about 0.80 kph
wet season troop),
season troop),
dry
the
differences
Symphalangus
were marked.
dry)
us
difference
1975),
did
allow
in
further
seasonal
moved slightly
(Chapter
90
of about
members
troop
The data
(Waser
would
productivity
The larger
than
for
albigena
The reduced
the reduced
dry > Anecdote's
seasonal
small
for
case
troops
distances
shorter
of
at Mt. Assirik
productivity
by foraging
of
than
further
travelled
lack
to be a result
1975) and Cercocebus
et al.
travel
baboons
This
be the
travel
significant.
the
the wet.
seems to
as
differences
in
to
wet > Fable's
statistically
that
is not likely
productivity,
(Chivers
was not
did
troops
wet > Anecdote's
the hypothesis
than
season
trend
larger
for
(Fable's
this
although
This
not
low productivity
to
was a trend
troops
smaller
to
but
troops,
further
in the wet season.
There
that
travel
habitat.
unproductive
of
to
ranges
may have been a response
troops
tended
Mt. Assirik
at
in
trees
in troop
they often
Combretum scrub
these habitats
5.26
therefore
that
at Mt.
sizes.
sat near the
on laterite.
The
tended to be earlier
those
than
for
This
8)
suggests
the
various
the
time
how
this
effect
in
the
time
This
could
be
of
to
feeding
the
sites
to
Having
feed
late
tended
may have
in
day,
and crossed
moved
to
be
the
16: 00hrs.
in
day
started
baboons
the
that
way.
they
clear
than
earlier
they
Similarly,
when
back
good
from
intervening
the
to
order
hunger
initial
their
satisfied
sites
the
and
the
of
a more leisurely
in
Assirik
suggests
beginning
the
at
not
Mt.
day,
at about
day
of
day,
is
it
but
at
the
of
occurred
time
rapidly
sleeping
rather
middle
generally
with
the
of
was bound to influence
baboons
of
parts
the
feeding.
and start
returning
in
speed
down, and began
slowed
late
(Chapter
and distribution
sites
The
noon.
after
habitats
were entered,
analysed.
and scrub-
scrub
on laterite
The location
ones.
habitats
and
been moving
food
find
various
of day, which
pattern
have
might
the
at which
forest
dry
to enter
productive
to the sleeping
feeding,
often
stationary,
time
were entering
relative
early
tended
10: 00, and grass
productive
habitats
at which
fastest
hottest
the baboons
less
than
earlier
the median
was after
were entered
that
The baboons
.
10: 00 while
or open woods before
grassland
habitat
any other
areas
rapidly.
baboon troops,
Most
do not alter
probably
including
their
ones
day range lengths
In some cases, for
changes.
the
in
reported
as daily
rather
is
than temperature.
by far
in
fact
watched
show no relationship
between day range
length
and
difference
is taken seperately
from both
the relationship
furthest
on cooler
troops
are
(their
pooled
may be spurious,
(a mean of 10.5 against
that
4, page 112).
4
a relationship
since
the troop
Figure
6.4km) also
and is enough to account
5.27
for
that
tended to be
days (means of 28.0°C and 29.9°C
respectively).
(1.9°C) is 9.5% of the range
of temperatures
were collected,
which
and
when data
tended to travel
(1970),
Saayman
and
from Stoltz
length
It
emerges, although
in productivity
between day range
when each troop
only
differences
wet season (Lee
correlation
temperature
is
of seasonal
The data
the best-documented
temperature,
further
example at Ruaha, baboons travelled
presumably,
thesis,
maximum temperature
.
each day in the hot dry season than they did in the cooler
pers comm), an effect,
this
This
over which the data
the observed correlation.
The observation
from any time
precision
temperatures
consistent
day ranges
successive
the maximum temperatures
to
be
or
speed
in
2) was
was not a
of
to
the
human
a
other
climatic
radiant
heat
(Gates
Index
If
movement.
1972)
there
was any
of
such
as
help
1976)
for
account
between
relationship
masked by the responses
was apparently
and geography
is
and Harrop
might
to
and there
variable,
(Russell
was
failed
deficit
Mt. Assirik,
load
season,
difference
this
pressure
any
the dry
in
especially
at
that
habitat
the
days,
between
the difference
greatly,
and vapour
of
1975),
et al.
speeds
troops
maximum temperature
troops
movement
and speed of movement it
climate
daily
To
humidity
suppose
(Chivers
changes
maximum
16: 00 (Chapter
15: 00 or
varied
degree.
one
Temperature-Humidity
the
the
of
to
reason
sunshine
that
about
successive
Temperature,
the
little
of
about
undetectable.
predict
the fact
with
some
with
of day range length.
good predictor
tended
that
the observation
be predicted
could
12:00, together
after
were not reached until
with
While
length
day range
that
the
through
area
which
of
they
were moving.
The distribution
Saayman 1970) or rainfall
in
sites
of the
on ranging
effects
were likely
together.
some suggestion
have influenced
the ranging
6)
since
the
speed
with
correlated
with
the
and
At Mt. Assirik
baboons.
season water
and of
they were normally
of water,
presence
of
the
sleeping
found close
although
superficial
there
water
is
may
of the baboons.
patterns
which
the
proportion
baboons
of
and may have been influenced
habitat
(Chapter
baboons
used
social
behaviour
presence
of riverine
for
troops
gallery
moved.
forest
In the
moved through
any given
in part
to be found in that
the
1976, Stoltz
Variables
The
which
for
here the effect
that
et al.
and Altmann 1970, Sigg and Stolba
of dry
location
to be confounded,
(Chapter
quadrat,
important
I have not analysed
5.9: Habitat
(Altmann
pools
may be extremely
press)
(Hamilton
of permanent water
habitat
present
in
was
that
by the number of food species
8) and in part
(Chapter
strongly
a quadrat
9).
by the areas which the
In
influenced
particular,
the
the
speed with
wet season the baboons engaged in long
5.28
periods
before
setting
in and near
behaviour
of social
the riverine
(Chapter
on the day's ranging
off
to
U.
they seldom showed these extended bouts of social
to riverine
ravines
often
spent
during
water
In the dry season
behaviour,
but returned
the day on most days.
in the
an hour or more sitting
The
woodland, and the
than the surrounding
degrees cooler
were several
troops
for
forest
gallery
forest
gallery
shade and grooming or
playing.
The ravines
the
containing
forest
gallery
laterite
to
habitat.
plateau"
was here
rain,
night's
This
forest.
laterite
behaviour
or
plateau
speed
trees
safety
in
for
on
from
the
off
and
dry
gallery
riverine
the effect
grass
which
in
troops
the
of
"grass
wet
could
the
of
movement
of
and moved out of
often
baboons
laterite
through
immediately
elsewhere
accounted
presumably
had on the
the
of
the
and the
absent,
metres
ran
were
The vegetation,
thin
a few
within
socialise
they
socialise
forest
left
when the baboons
In the morning
plateaux.
gallery
riverine
the
on
wet
season.
Few food species
grew in scrub and scrub
the baboons moved rapidly
provided
through
many food species,
quadrat
speculate
habitats
that
in
(Chapter
food
8),
habitats
far
variance
accounted
gallery
provided
other
than it
attractive
much of the
In the
dry
speeds in
their
altered
this
analysis
suggests
that
has shown.
there
season,
the
the local
with which the baboons entered
habitat
forest
types
found
seemed to
because of the quantity
in
the
be attractive
factors
the habitat.
depended on the
quadrats
quadrats.
Forest
to
the
of food to be found there
5.29
The small
the most important
of food within
concentration
different
were other
the speed of movement of the troops,
The frequency
possibly
less
bamboo shoots.
than
by habitat
for
of which was probably
of
in
enough food
found
troop
probably
more sensitively
influenced
riverine
contained,
especially
Baboons at Mt. Assirik
proportion
I
was somewhat less productive.
however, dry forest
which
but only in the dry season.
In the wet season dry forest
was in the dry season.
troop's
a
moved more slowly
open woodland to become relatively
for
hand,
through
troop
and Fable's
the wet season the
grassland
8), and
Open woodland, on the other
them.
the more open woodland it
(Chapter
them,
(Chapter
and
first
8) and
trees.
the second as a source of water and sleeping
As predicted,
how frequently
results
speed
with
sensitive
to
between
troops
While
baboons
to
in
the
and may
troop
and
density.
In
was rare
off.
They also
tended
to move more slowly
was higher,
distance
that
the probability
the inverse
This
finding
way.
time in quadrats
understanding
environment
of
the
nearest
the further
of
the
distance
may therefore
benefit
likely
to
be
a given
to that
with
from
5.30
baboons
sleeping
research
at
any
sleeping
site.
of Sigg and Stolba
their
environment
one or more
the baboons spent
so that
which
the
decreased
quadrat
on cliffs
slept
to
distance.
with
they were from the sleeping
found at Mt. Assirik,
had been
baboons
from the nearest
contrast
waterholes,
in
the
distance
linearly
decreased
P. hamadryas did not exploit
manner
in
grass
the
finding
However, the baboons they studied
from
of
trees
were
areas
by the
they would enter
in direct
who showed that
these
areas
where the visual
in areas
influenced
site
square of their
is apparently
kilometres
together,
a sleeping
the
after
the
good visibility.
of very
The probability
site.
from
with
that
were strongly
patterns
sleeping
in this
which
than were areas
more productive
Ranging
may indicate
from
3
in
predicted
far
poor
Chapter
entered
see a troop
a plateau
in
tends
of
areas
as
season,
crossed
This
given
preferentially
to
between
negative.
avoided
which
with
correlation
arguments
dry
the
apparently
It
the
to
seldom
press)
adjust
difference
this
frequency
the
baboons
the
season,
Similarly,
to
was apparently
for
on the
and they
given
temperatures
failed
troop
was strongly
support
size.
baboons
higher
nearest
troop
season,
wet
that
lend
,-
the
density
These
to higher
reasons
effects
density
hypothesis
the
visual
in
and visual
entry
predation
burnt
shade.
to
Fable's
The
seasons.
equivocal
quadrats
introduction,
(in
both
the wet
regard
Anecdote's
while
shade,
in
while
is not clear.
of
support
dry
to
respect
entered
visibility,
on
cooler
shade
However,
areas.
shade had rather
frequency
without
deciding
in
part
season,
the baboons may have responded
the
out
dry
the
ranged
apparently
that
suggest
by seeking
its
troop
in
was entered
a quadrat
Fable's
season
shade
an important
played
apparently
sites
Further
cliffs.
exploit
the
and water
on refuging
more
sort
of
normally
animals
(eg
and Vehrencamp 1916b, Covich
1978, Bradbury
Andersson
1976, Hamilton
and
Watt 1970).
types
Topographic
(Chapter
8)
typically
moved through
in
areas
which
the
troops
P. hamadryas
described
for
travelled
through
once
the
always)
(Chapter
returned
6),
was related
device
several
area
to avoid
by
used
food
items
types
which
behaviour
were
few
contain
topographic
whereas
they
times
to
the
that
foraging,
reinfection
sleeping
this
revisited
a
lay
Sigg
systematically.
I believe
to
at
In general,
study.
the
to exploit
days
Some quadrats
species.
in
for
repeatedly
particular
to
for
social
slowest.
On some occasions
sometimes
fastest,
were moved through
occurred
likely
were
which
on
during
stretch
and Stolba
a week.
Other
however,
it
Since
that
site
systematic
troops
coverage
as suggested
of
like
the
of
left
their
that
tended
not
morning
home ranges
by Freeland
of the troop.
only
(but
frequently
had
be
and might
troops
a
those
were visited
quadrats
they
the pathogens
5.31
press),
seemed that
the
fruiting
routes,
(in
days,
successive
the
on well-defined
and was not,
with
areas
(1976)
a
CHAPTER 6: Sleeping
Sites
6.1: Sleeping
Introduction
Sites:
The presence
baboons
not
(1979)
are
in
sites
or
present
in
1975)
1971).
In
the
15 (Altmann
sleeping
some are
usually
and Hall
1965),
sleeping
trees
reported
to
and in
and
baboon
in
sleep
on
others
baboons
only
to
1973; P. papio,
Bert
in
sleep
et
been
on cliffs,
in
Harding
1967a,
al.
20% of
only
trees
1964,
Rowell
so that
than
by fewer
and T. gelada
known
are used,
preferred
are
those
1970, DeVore
has
(P. anubis,
others
and Keith
are
that
sites
for
is
that
while
each,
P. cynocephalus
and P. hamadryas
of
in
Although
found
and Altmann
some trees
troop
baboon
(1980)
had one site
may be accounted
1971).
used by a troop
sites
(Altmann
them
(Jouventin
Deag and Crook
Of the
al.
et
to
sphinx
Müller
had several.
1951).
trees
cliffs
The number of
groves,
nights
species
Stoltz
P. ursinus,
sleeping
(Lumsden
three
other
over
preferred
available
ten
about
Bert
used by a single
sites
a waterfall
of
are
Mandrillus
availability.
downstream
gorge
70% of
more than
of
upstream
in the
sleeping
their
sites
of
sleeping
of
number
(eg
sites
from
ranges
use one to
(eg Macaca sylvanus,
1970).
and Altmann
a function
troops
macaques
by the
normally
suitable
differ
they
this
home
sleeping
number of known sleeping
The greatest
probably
traditional
when many apparently
and some of
sites
1979)
the
of
or
Rasmussen
1961).
determined
Baboon troops
home ranges.
whether
and DeVore
quality
relative
may be largely
(Rasmussen
even
(Washburn
an area
may determine
sites
sleeping
of
the
that
their
(Ransom
the
absence
troops
twelve
1967b)
is
or
believes
neighbouring
and Water
some
1976;
Emory pers
comm.
Some primates may respond to the threat
small
dispersed
Hylobates
al.
Nathan
Chivers
concolor,
1965),
sharing
groups
while
sleeping
(eg Galago
1972, Erythrocebus
rely
with
groups
sites
other
1972, Kummer 1968a, Kummer and Kurt
in
group
tend
to
cliff
face
(Dunbar and Nathan
sleep
of predation
subgroups
within
and Doyle
Bearder
senegalensis,
baboons apparently
by sleeping
Hall
pates,
on numbers, with
(Bert
et al.
1963).
a single
et
often
1967b, Dunbar and
The baboons in
grove
each
or on a single
1972, Kummer 1968a, Kummer and Kurt
6.1
1974,
1965a, Hall
groups
in
1963,
6.1:
Map
'77-'78
SLEEPING SITES IN VICINITY OF Mt ASSIRIK
Xanadu
Mt Assirik
Elephant Rib
Assirik Valley "
*
wild
Dog
Flycatcher Falls'f"
Fork
North Cliffs
Waterfall
Old Camp,, Camp
Stella's
Valley
(West)
CrossValley
Secretk
Valley
Key:
0Sweet
Spring
f Usecommon (but possibly seasonal)
0 Userare or unconfirmed
I km
1980), thus
Muller
in the safest
sleep
apparently
some further
giving
faces
on sheer cliff
to them (DeVore and Hall
available
(Bert
trees
They
et al.
1967b, Nagel 1973) or
(Kummer 1968a) even when trees
(Booth
1956a, 1956b, Hamilton
1976, Nagel
et al.
1973).
are available
No quantified
data are
baboons might use to choose one possible
on what criteria
available
places
high in the highest
1965), either
from predation.
protection
roost
over another.
move after
dark.
a sleeping
site
Baboons rarely
was displaced
journey
from
11 km from
of
hours
of daylight
to
from
chase
wild
dogs
One troop
of
baboons
(Davidge
sunrise
Macaca mulatta,
by another
habitually
grove
left
Baboons
at
pens
50 minutes
cliffs
differ
hundred
A
have been seen
(Lee
nightfall
may therefore
1971c).
has been observed
site
and baboons
sleeping
may move several
which
a second
a troop
after
(Saayman
troop
(Kummer 1968a)
a sleeping
1977).
to
site
an occupied
in the closing
One such move occurred
at
before
troops
from
metres
comm).
of
(Vessey
night
1973).
The location
of sleeping
of baboons (Chapter
whether a troop
(Hall
new site
6.2: Sleeping
at
sleeping
One troop
to
close
cases
used
closest
several
sleeping
in
had at
least
first
troop.
One of
was a single
summit
of
the
left
in
slept
field
on
morning or moves to a
that
trees,
permanent
laterite
made of
8 well-separated
1.4
sites
6 in
its
Ceiba
(Map
home range,
sites
6.1)
was only
five
have been
few ledges.
and were in
5m, range
0-35m).
(median
distance
between
some of
which
which
metres.
contained
The other
were common to
used once to my knowledge.
pentandra
growing
was ringed,
about
6.2
and having
hundred
of
although
may not
distance
sites
by several
separated
these
Mt. Assirik
km)
on cliffs,
grew in ravines
(median
water
not
These cliffs
area.
being
sites,
groves,
troop
the
Mt. Assirik
least
at
neighbouring
"site"
it
that
they chose to sleep in normally
The trees
most
depending
Results
Sites:
for
on the ranging
1962a).
were many cliffs
suitable
influence
For example, day range may differ
to the site
returns
The baboons
there
5).
has a major
sites
below
the
2 km from
cliffs
the
with
nearest
the
This
which
major
0
Its
site.
use as a sleeping
suggests
that
been of
great
there
may have been other
importance
14,10
and 7 trees
trees
came from
three
of
data
collected
or
survey
the
taller,
but
trees
cordifolia,
Erythrophloem
or taller.
The median height
25m, principally
over
the other
species
in
sleeping
the
and
reached
Cola
senegalensis,
species
7
SAPP team
vines)
pentandra
(Fisher
slept
(21,
25 m
reached
in by the baboons was over
slept
chose Ceiba
they
sites
africana,
ravines,
Khaya
whose
The remaining
(excluding
tree
of
of the trees
which
the
and two unknown
the baboons
in
and of
Afzelia
pentandra,
suaveolens
because
which may have
Mt. Assirik,
at
cordifolia,
growing
Ceiba
only
the wet season
sites
censused in four
21 species
that
showing
little-used
baboons
(Cola
species
In
10 baboons in
44 were Ceiba pentandra.
respectively),
suaveolens).
a
the
Of 52 trees
Erythrophloem
15m
to
I am not aware.
existence
by about
site
preferentially
one-tailed,
exact,
p=
0.003) (Table 6.1).
Table 6.1:
Dimensions
nS
Species
(metres)
of Trees used as Dormitories
by Baboons
Height
of Tree
Height
of first
Branch
Mean
Mean sd
Mean sd
Mean sd
Mean sd
sd
Distance
Distance
to Water to nearest
Dormitory
Diameter
at lm
Ceiba
44 8
27.0±6.7
16.1±5.7
1.710.9
7.6±7.9
10.8±9.4
32
16.7±2.9
8.0±0.0
0.7±0.3
3.3±1.5
7.7±2.5
suaveolens
45
20.5±8.4
4.6±3.9
0.9±0.4
3.5±1.3
9.8±4.5
Afzelia
africana
12
25.0
6.0
0.5
1.5
pentandra
Cola
cordifolia
Erythrophloem
KEY:
found
to
number
s=
number found in line
routes
-
10.0
-
-
be used as dormitories
n=
Access
-
transects
into
the
sleeping
one
representative
pentandra
having
only
apparently
climbed
up the trunk
trees
of the tree
6.3
of riverine
were variable,
into
itself
forest
gallery
which
(Table
with
the
6.2)
Ceiba
baboons
Table 6.2:
Access Routes into
Species
Sleeping
Trees
Access Route
other tree(s)
Number
vines
self
Ceiba
63
54
8
1
Cola
cordifolia
3
0
1
2
Erythrophloem
suaveolens
4
0
1
3
Afzelia
afrieana
2101
pentandra
Fable's
on u of
morning
(chi
site
however,
site
troop
full
the
was equally
the
day ranges,
= 9.75;
squared
df
likely
it
on 19 it
while
= 1;
to
that
site
left
moved to
(12)
previous
a different
Anecdote's
p<0.005).
return
the
as
to
troop,
a different
move to
(14).
The troops
there
to
returned
were never known to move away from a site
was commonly much activity,
bursts
of
loud
with
including
vocalisations,
screams and geks (Ransom 1971),
defaecation.
copulation
Copulation
grunts
observer,
by
Occasionally
leopard
vocalisations
Silent
periods
nights
and after
calls,
by
or
would
during
nights.
the
and before
6. u
sound
and
nights
and
Outbursts
first
and of longer
light.
with
urination
moonlit
of
but
weaning
of
triggered
distant
accompany the wind preceding
seemed to be more frequent
midnight
roargrunts,
but were readily
spontaneous,
interspersed
some squabbling,
observed
on other
periods
wahoos,
and with
was
were heard
were sometimes apparently
silent
at night,
duration
calling
by the
baboons.
a storm.
on dark
6.3:
Sleeping
Ceiba
within
ideal
they
were
singled
was
that
a
the
usefulness
or
6.4:
Water:
the
food
absence of
et
1976; Hall
sleeping
1963b).
picked
their
their
roosts,
trees
hardly
used
way
that
or
surfaces
tree
it
and
a tree
gingerly
they
so that
smooth,
giving
were
surprising
The upper
through
against
like
water,
or
one
along
the
along
mute evidence
of
predation.
limits
1973,
Stolba
any troop
day (Altmann
1973, Stoltz
and Saayman 1970 p116).
been studied
obtain
Blake
1971, Collins
Kummer 1968a,
their
in
waterholes
Stolba
1979,
known to
are
Hamilton
et al.
provided
artificially
prep,
both
livestock
6.5
is
1966).
freely
probably
from
and
DeVore
fresh-water
Some populations
(Harding
1976,
The
available,
drink
every
day,
1970 p140; Nagel
comm, Nagel
and Altmann
et
away from water
that
1973,
1970,
1961),
lakes
have
(Aldrich-
season from rivers
Lee pers
Washburn
drink
(Hamilton
Most baboon populations
(Altmann
1976, Ransom 1971).
for
influencing
(Hall
and Altmann
in the dry
water
or
Thus
behaviour
drinkers
1971),
baboons
1976, Kummer 1968a,
et al.
When water
of
or
1961).
has been known to stay
(Lindburg
other
every
1973)
of baboons,
1979) and social
1976).
al.
et
Macaca mulatta
et al.
Washburn and DeVore
Hamilton
presence
distribution
the
in the ecology
1970,
the
as much as
Baboons are obligate
that
occasionally,
populations
is
dormitory
moving
sites,
and Altmann
(Hamilton
5 days
and Keith
it
These
trees.
Baboons
of
Nagel
in press,
interval
baboons,
or,
absence
1976,
al.
(Altmann
longest
is
or
or
Sigg and Stolba
al.
tree.
were worn
as a defence
is of major importance
ranging
horizontally,
its
Introduction
The presence
water
thorns
nearly
a traditional
from
or
out
frequently
most
C. pentandra
used
moved to
covering
and
as sleeping
used.
rarely
thorns
grow
baboons
were
between
never
was
of
the
a dormitory
with
a 25 m tall
of
baboons
distinguish
as they
(Hamilton
trees
by the
of
that
top
that
was rarely
branches
the
of
tree
branches
the
out
to
branch
Its
for
were
easy
buttressed
a
protection
branches
the
5m
only
that
of
is
bark.
smooth
probably
Discussion
pentandra
otherwise
often
Sites:
Hall
though
(Hall
Stoltz
1966,
three
1963a,
make use of water
Stoltz
and Saayman
1970).
At some sites,
(eg
year
Hamilton
in
season
which
Okavango).
et
water
in
the
Blake
1969
holes
made by other
troop
in
(Cheney
et
for
provided
Troop
, to
(Harding
1970,
size
permanent
sources
further
from
found
(data
p<0.001)
of
rain
in
from
interest,
a year
and "by the
but...
water,
its
may dig
similar
One
fight
to
seen
it
and
is
aethiops
during
of
times
water
the
greater
relief,
baboon
to
by the
location
with
smaller
than
larger
and Keith
surface
ones
since
(mean
end of
1977).
in
station
park
is
dry
season,
it
Most of
the
region
does not flood,
6.6
press).
relative
29)
(t
being
= 9.9;
1973).
season the
the
in
41)
size
the GPN de NK is
as few as 57 days receive
rainy
water
(mean size
troops
may
(Altmann
home range
the
of
sources
and usage
Stolba
and
near
previously
areas
shape,
size,
Sigg
Euphorbia
to
water
into
spread
1976,
"more
were
Uganda was high
in
of
centre
permanent
(1975)
Artificial
absence
al.
of
damage
Park
Home range
or
estimated
source
al.
et
between baboons and water
end of the
at
baboons
et
Stoltz
Boese
161 troops
Of
water.
1972).
at a nearby meteorological
(Dupuy and Verschuren
with
1976
Cercopithecus
a known
and
(Lock
water,
water
The relationship
particular
no
comm).
1976),
19 had the
National
presence
may be influenced
by
water,
water
Hamilton
but
all
seen
1976).
on the
from
5 km from
have allowed
cattle
Altmann
and
than
from
6km from
depend
therefore
been
al.
may die
far
(1973),
24 troops
walk"
by them
unused
less
the
and absent
water
et
low-ranking
travel
the Queen Elizabeth
in
candelabrum
like
rarely
and Keith
hour's
an
water
is
and Aldrich-
Lee pers
has
(Hamilton
baboons
and maintain
1976,
et al.
Wrangham 1981),
apparently
None of
than
enlarge
the
et al.
(Crook
water
Namib desert
the
baboons,
home ranges
water.
in
to
press,
by Stoltz
their
of
clear
or
foul,
or
of
there
others
(eg Hamilton
critical
obtain
some months
1963a).
Baboons
censused
Canyon
access
in
(Hall
shortage
to
in
at
unavailable
(Hamilton
animals
for
al.
is
water
low-ranking
that
likely
becomes
riverbeds
Kuiseb
troop
while
Kummer 19684 p164),
p 214,
the
Kuiseb),
shortage
sand of
may be severe
stress
1976
al.
When surface
holes
another
water
but strong
of
more than
1mm of
(McGrew et al.
1981),
almost
is
entirely
under
like
semi-desert"
around
Mt. Assirik,
seasonality
is still
(Chapter
apparent
6.4.1:
Sleeping
The
2).
or
presence
powerful
determinant
sleeping
sites
water,
that
and
this
Unfortunately
6.4.2:
the
choice
in
turn
six
more
than
(Chapter
the
cause
live
the
ranges
this
of
each
metres.
to
troops
was
presence
or
P. ursinus,
of
the
of
important
troop.
statement.
in
in
distances
each
which
All
of
these
and
yet
the
wet
(Chapter
troops
of
fragment,
particularly
large
Furthermore,
was separated
features
access
from
the
large
7)
in
to
at
next
by
have
might
when the
season,
had
displayed
they
in
(Chapter
a day
troop
3)
tended
a remarkable
visibility
was at
worst.
P. hamadryas
tended
coalesce
temporarily
in
waterhole.
This
rendez-vous
at the start
certainty
the
whereabouts
of
the
6.4.3:
permanent
(File
in
three
during
usually
afternoon,
by
prearranged
water
the
(Stolba
ranging
sources
for
the
morning
near
troop
a
with
1979).
and
good
some
A Knowledge
may have been important
but also,
survival,
the dynamics of troop
if
they
structure.
and Water
many of
intermediate
fragment
not only
for
At Mt. Assirik
mansonii
early
of the day's
rendez-vous,
Parasites
to
is
the baboons at Mt. Assirik
were used for
to
in
long
2).
sites
a thousand
Bands of
for
a troop
two
water
surface
for
that
a
to permanent
near
that
day
support
found
noted
sites
the
in
when
p113)
influenced
travelled
and
and
sleeping
cohesiveness,
of
season,
sleeping
no data
give
being
season,
may be
sites
Kummer (1968a)
dry
wet
of
Mt. Assirik
at
visibility
least
its
the
sleeping
Water and Group Cohesion
home ranges,
to
the
in
near
use.
and Saayman (1970,
they
Baboons
poor
in
abandoned
influenced
water
water
home range
were used
Stoltz
widespread.
of
absence
of
that
were
Home Range, and Water
Sites,
et al.
host,
genera,
the
of
Schistosomiasis
prep).
any one of
all
baboons were infested
17 species
which
live
6.7
in
of
is
Schistosoma
with
transmitted
pulmonate
marshy areas
snails
in
to the
belonging
slow-flowing
in
water,
the
Infection
faeces
and
urine
occurs when contaminated
If,
Altmann
as
to
the
strongly
adapted
"contact
between
lips
expected
to
drinking
avoid
and
when there
schistosomes
(1970)
avoidance
from
was
sources
in
the
field
of
water,
by
likely
available
were
so
that
schistosomes
baboons
minimised,
sources
were
1968).
drinking
suggest,
infection
of
water"
(Barnes
animals
is drunk.
water
Altmann
and
infected
of
be
to
which
be
would
by
contaminated
were
less
likely
to
be contaminated.
6.5:
Water:
Methods
Water
sources
or
presence
flowing.
Only
absence
The turbidity,
one water
The
and
myself).
-
were
seen,
which
water
if
present,
and extent
was monitored
state
of
showing
of
sources
whether
still
riverbeds
noting
or
not
and permanent
was
Harrison
they
whenever
them by the
to
it
was recorded.
McGrew,
recorded
the
baboons.
A
or semi-permanent
sources.
The water
drinking
there
could
surface
inferred
was
disappearance
by
the
be seen by the
always
attitude
baboon into
the
of
not
of
the
a depression
in
observer,
baboon
the
or
ground
and
by
in
the
which
was water.
Sources at which baboons drank were defined
by
infestation
first
area
unassociated
pan).
Sites
site
infestation,
beneath
last
sleeping
stream
or marshy
area,
with
with
"uncontaminated"
sleeping
low
a
site
but
site
(ii)
of
a sleeping
or
surrounded
reedbed
was closer,
infestation
of
site
or
(ii)
upstream
(iii)
and
pool
are
than
a source
referred
of
the
of
the
in
to
an
unassociated
as
risk
of
located
1km downstream
a reedbed.
of
on a laterite
as having
a high
if they were (i)
and less
from
a low risk
tkm downstream
(eg a rainwater
by or draining
6.8
(i)
than
more
whichever
sites
risk
as having
located
were defined
to as "contaminated",
and referred
or downstream
were
Sources
sites.
sleeping
they
on the
site
sleeping
if
schistosomes
sleeping
last
the
by
water
(by
was
when I was led
major
monitored
systematically
other
was usually
map was prepared
were
and,
depth,
source
area
with
of
a
xan 6.21
--
----
km
qn1
a
P
fý_
te
kc
Cm
,
lb
./u
h
'76/7,77/8
"
dry
Key.. water throughout
seasonsof
0 dry before start of rains
dry
o water at start of
season
a
b
c
Elephant
Rib
Assirik
Valley
Poacher Bend
d
"
f
Lower Middle Lion Valley
Upper Middle Lion Valley
Head of the Waters
y
h
Cross Valley
Secret Valley
L
j
k
s
Snaking Rock
Fork
Tributary
Sam's Kapok
Three Tribs
n
Lower Elephant
o
Xanadu
p
q
r
s
t
u
Baboon Spring
Fork Falls
South Plateau Waterhole
Dead Kapok Rapids
Flycatcher
Falls
Whaleback Wallow
Rib
6.6:
Water:
Results
The wet season lasted
of the wet season,
from about June to September,
in storms at night
a week, typically
half
an hour.
the
In this
plateaux
in
In the middle
fell
rain
lying
was abundant,
to
rivulets
the
small
times
several
in which 50mm or more might
season water
and running
which flowed
from May to October.
roughly
fall
in
in puddles
on
(Map 6.2),
streams
sometimes more than a metre deep.
continuously,
Table 6.3:
Nature
and Seasonality
of Water
Sources
Season
Mid End
Mid End
Wet Wet
Dry Dry
Nature
Source
of
Aug Nov
Rain water puddles
Running water on plateaux
Stagnant
water on plateaux
Running water in deep ravines
Stagnant
In
the
to
restricted
March
1977 the
pools
at
the
season
of
contaminated
with
faeces
in the trees
overhanging
In
35m of
but
all
the
(Flycatcher
streambed
water
in
from
the
field
water
waa
and Table
6.4).
In
primates,
lay
area
The pools
vegetation,
the
February
were
in
many
baboons,
mostly
less
all
in
and
stagnant
that
than
cases,
slept
the water.
one known cases
choice
Falls),
(Map 6.2
ravines.
rotting
by
and
Home Range, and Water
a streambed.
affected
the
Sites,
fell,
area
sheltered
with
++
field
remaining
bottom
Sleeping
++
the
only
filled
6.6.1:
+
rain
in
(f)
+
no
8 sites
deep,
5cm
+
water in deep ravines
dry
Feb May
The
presence
of
sleeping
near
contained
(n=239).
a seasonal
water
(chi
6.9
of
baboons
water
site,
stream,
squared
in
since
the
stream
one
was rarely
6.46;
=
in
slept
df
trees
within
apparently
sleeping
occupied
= 1;
site
unless
p(0.02).
6.4:
Number of Observations
Table
of Drinking
(See Map 1 for
Name of Source
Elephant
Water
Sources
Locations)
Season
Wet Dry
Visibility
Availability
Mid Dry End Dry
Rib
1
0
Poor
Valley
4
5
+
Excellent
+
1
0
1
0
1
3
Good
Poor
Poor
+
+
Head of the Waters
Cross Valley
Secret Valley
Snaking Rock
9
5
0
4
7
9
0
3
+
+
+
+
+
+
Good
Excellent
Poor
Excellent
Tributary
Fork
Sam's Kapok
Three Tribs
Lower Elephant
Xanadu
3
3
+
+
+
-
0
2
0
1
1
0
0
0
Poor
Poor
Good
Excellent
Good
+
+
+
+
+
+
+
+
?
0
0
0
0
0
9
0
0
0
1
0
0
Good
Poor
Poor
Poor
Poor
Variable
?
?
-
Assirik
Poacher Bend
Lower Middle Lion
Upper Middle Lion
Valley
Valley
Rib
Baboon Spring
Fork Falls
South Plateau Waterhole
Dead Kapok Rapids
Flycatcher
Falls
Puddles, Rivulets,
etc.
Range use
sleeping
for
at Various
of
There
occasions.
middle
The troop
of
the
the baboons'
dry
of its
season.
when water
in the middle
subsequently
was no surface
was no trace
-
was present
1 was led to this
time on 14/7/78,
study.
and there
changed
(Xanadu, Map 6.2).
site
the first
months
may have
-
site
at
another
by the baboons
of the wet season after
this
visited
site
on
several
the
dry
season,
this
site
in
use as a sleeping
site
on 22/1/78,
water
The areas
at
in
the
north-western
home range were only used when this
sleeping
12
in the
portion
of
was known
site
to have water.
Baboons were seen to drink
and 33 in
to drink
the dry.
baboons
They drank from 14 sources,
from 6 only
On 8 out
to
of
one of
on 73 occasions,
40 in
the
sites
on which
used
6.10
in
I
both
wet season
of which they were seen
in the wet season and 3 only
12 occasions
the
followed
seasons,
in the dry.
fewer
than
baboons
about
were
20
either
already there or one or more groups or baboons arrived
these 12 cases, all
a cohesive group.
2 (or possibly
later.
the baboons present at the site apparently
In 9 out of
moved oft
in
In two of the remaining cases the group fragmented into
more) parts, while in the other case, I lost
the baboons in
the time taken to climb out of the ravine.
6.1.1: Drinking
Over
the
with Time of Day
whole
baboons were seen to
year,
drink
more frequently
between 12:00 and 14:00 and between 16:00 and 18:00 than they were between
14: 00 and 16:00 (chi
6.5).
No other
day was usually
Assirik
a 5.55; df s 1; p(O. 05 In both
squared
differences
significant
were noted.
The hottest
between 14:00 and 16:00, which suggests
their
did not regulate
drinking
with
cases)
that
(Table
time
of
baboons at ft.
the heat of the day, but drank
in the middle and at the end of the day's journey.
Table b. 5:
drink
to
Baboons
Frequency
atý
with
aeon
which
were
. rriý.
_-_.
.. `rar
"
various
Time
Times
of
Ses5on
Day
Year
Wet Dry
6,1,2:
_Parasites
The trees in
distance
06: 00-07: 59
6
6
12
08: 00-09: 59
7
2
9
10: 00-11: 59
12: 00-13: 59
14: 00-15: 59
3
8
1
4
6
3
7
14
4
16: 00-17: 59
10
4
14
18:00-19: 59
5
8
13
and Water
which
the
5m, range 0-35© )*
about 50-75% of aaaaa.
high probability
that
would be found
in
if
it
baboons slept
and their
grow close
6.11
in
that was sampled there was a
the baboons had slept
the corning,
(median
branches overhung the water
At the only site
in
to water
above the stream, fasces
goody or
marshy areas were
intermittently
present
invertebrate
formed
Conditions
Sites
was
which
of
the
site;
at
this
which
therefore
right
streams,
and while
the
from
dry
that
seasons
general,
baboons did
of
Marks
Waters".
some underground
source.
the
above
often
was present
in the vicinity
defaecate
at
to
avoid
of
directly
into
waterholes
into
trickled
occasions
this
of
and 1977-78.
bordering
on rocks
for
ground
upstream
Water
to
seem deliberately
not
the
of
existed
1976-77
observed
had on several
urine
sites
site.
present.
present
seen to defaecate,
trees.
evidence
from
not
were
of
also
50m downstream.
They were never
source.
except
was
one was about
transmission
named "Head
source
No sleeping
year.
Schistosoma
of
the
be contaminated
emerged
water
hosts
for
the
they may have
downstream of a sleeping
to
water
Baboons were frequently
water
intermediate
a water
shows
the
of
throughout
a water
larger
the
likely
spring
nearest
site
less
Map 6.2
months
several
for
were
were
a site
Upstream
the
of
to any baboon drinking
example,
this
habitats
suitable
schistosomes
This
all
fauna of these reedbeds was not investigated,
mansonii.
For
along
gave
water.
In
contaminating
the
water.
Although
(chi
squared = 12.25;
sources
in
were
contaminated
drink
baboons
ones,
apparently
avoided
df = 1; p<0.001)
general
so that
from contaminated
for
easier
the
sources
drinking
(Table
the
occasions
contaminated
6.6),
the uncontaminated
to
observer
on which
water
than
see
were
baboons were seen to
are significant.
Table 6.6:
Frequency with which
or less likely
Probability
Contamination
Season
Wet
Dry
of
Low
High
*includes
Furthermore,
Baboons were seen to drink
at Sources
to be Contaminated
with Schistosomes
32*
10
Total
23
8
n of
major
55
18
more
sites
7+
8
9 observations
from uncontaminated
of baboons drinking
in the Wet Season
rainpools
and rivulets
the
baboons
tended
to
spend
6.12
longer
drinking
from
sources
may have been contaminated
which
(Table
than
they
did
at
puddles
or
rivulets
6.7).
Table 6.7:
(sec) for which Baboons
from Different
Sources
Duration
Drank
From Puddle
From Stream
2
12
5
23
6
6
6
27
27
27
8
30
8
31
9
32
10
38
13
13
38
Mann-Whitney
U Test
U=1
Two-tailed
p<
6.2:
Discussion
Water:
At
Assirik,
Mt.
In
regimes.
the
baboons
experienced
water
was abundant
the wet season,
baboons were seldom more than
In the dry season,
permanent water
Altmann
to
P. hamadryas groups
in
a dry
sources
result
baboons.
and Altmann
matter
few
as
studied
that
I
shortage
Such an arid
was
animal
than
year
that
waste.
water
of
14 in
6.13
about 8
40 krýr/aq for
contaminated
Although
neither
sources
as
which had only
might
frequent,
water.
80 km,Aq
13 in
become a serious
may be fairly
obtained;
with
1977-78 several
as "permanent"
could
distribution
and
water
from surface
everywhere
permanent
have described
water
1970,
by Kummer (1968a),
season more severe
which
and
or
and readily
of metres
in its
different
two
q (compared with
100 km
1979),
vegetable
restricted
a few tens
became limited
in
Stolba
P. hamadryas;
decomposing
water
sources
P. cynocephalus;
was
.
002
of
dry
for
with
troop
were
the
2-4 sites,
the
water
up,
with
the
problem
for
the
given
that
the
annual
may differ
rainfall
between years
by nearly
50% (McGrew at al.
1981).
Reduction
baboons
to
in
survive
replenishment
the
all
in
on water
of
If
this
the
North
6.2.1:
Sleeping
There
the
an area
that
use of
which
These data
in
range
in
size
encompassing
postulated
that
to small
living
in
arid
water
but
sites
troops
at
the
access to water
used
that
the
permitted
the
season,
dry
a reasonable
limited
when
by
Indians
water
evidence
baboons
to
9
home ranges
the
to an observed
holes
from
to
permanent
waterholes
$8 "major
and
1970).
Although
by the availability
the small
of water,
of permanent
this
range on the basis
the home ranges
of
area
by the
arid
areas may be restricted
Gombe Stream
and their
of
their
National
home ranges
6.14
many troops
of
Park
in
in
distribution
For
environment.
Tanzania
were probably
of
baboon troops
in
less
they
known home range was due
initial
determined
their
home
season home range
a few hundred metres
then,
increase
season
home ranges
aspects
of
or absence
18-fold
a dry
a wet
Altmann
and
they
in
range
was disused.
and the
troops
was
Further
by the presence
governed
were within
since
areas,
by other
were
for
slow
Shoshonean
of
site
water
In general,
of
part
size
P. cynocephalus,
of
on
season may be restricted
The home ranges
was in
troops;
sites.
(Altmann
seems likely
work.
in
when the
was not applicable
sample size,
one month's
be too
thirst
the
allow
large
well
was present.
home range was restricted
"new" sleeping
It
in
sleeping
touching
or
rainpools"
water.
that
a troop
temporary
the
entered
on 5 permanent
centred
sites
site
were in part
explanation
range
group
sleeping
a sleeping
suggest
near potential
This
size
might
support
might
their
group
when no water
was not
baboons at Mt. Assirik
of water
water
limit
to
to
meagre
satisfy
then
season
Home Range, and Water
and disused
that
suggests
to
dry
1955).
evidence
nearby
present
into
Sites,
is
troop
case,
(Steward
America
subsurface
thought
same circumstances
the
too
supplies
a large
were
in
size
from
a pool
members of
time.
the
mean troop
of
the dry
water.
by
not
example,
the
had "unlimited"
constrained
by the
4
presence
of
difficult
to
interpret
the
and any independent
size
(Ransom 1971).
troops
neighbouring
results
of
6.2.2:
Water Sources and Group Cohesion
dry
foraging
group
problem,
but
permanent
water
was
wet
season,
baboons
a given
1961)
in
of
wandering
troop
(Ransom
least
at
In
a
study
that
the
dry
season
bands
stay
together
after
they
assembly
was
troop
P. anubis
of
scattered
widely
the
not
have
the
P. hamadryas
the
baboons
6.2.3:
There
other
the
arid
reassembly
troops
as
studied
to
at
the
did
that
Washburn
on leaving
(1968a
p12)
reassemble
end of
of
at
(1971)
the
wet
the
day's
(in
did
not
such
that
reported
and
sleeping
a
became
site
journey
mid-journey
and Stolba
found
season
parties
near
the
but
water,
the
into
split
that
predictable
Kummer
al.
et
the
poor,
difficulty
In
by Sigg
(eg
not
without
water.
that
were
troop
midday
the
water
and
at
region
when there
in
does not
assembly
of
that
of
press)
or
at Mt. Assirik.
Parasites
sources
an
same significance
the
left
foraging,
while
This
evening.
in
regular
find
came together
units
generally
also
P. hamadryas
of
foraging
together.
so
always
used
troop
these
probability
was
were
the
a
When the
even
That
at
high
Aldrich-Blake
seen.
source,
together
visibility
could
water
move off
troop
one
members
1971).
would
separated
been
may have
cohesion.
sources.
by the
where
1977,
and mean
have
may not
the
area
drawn
good,
Mt. Assirik
at
permanent
was suggested
group
cohesion
troop
temporary
many
Gombe Stream,
for
catchment
troops,
the whole
movements
focii
associated
different
source,
At
the
water
of
and DeVore
the
group
the
at
surface
members
smaller,
as
home range
and Harvey
was relatively
was
in
reassembled
when visibility
sources
be
1970).
et al.
size
in
foraging
was
season,
between
(eg Clutton-Brock
variable
and May 1976, Turner
the
may therefore
correlations
Milton
In
It
is
likely
parasites.
and Water
clear
to
evidence
from
be contaminated
The baboons of
Mt. Assirik
with
Amboseli
6.15
that
schistosomes,
might
baboons
and,
drink
at
possibly,
have been subject
to
different
dealing
with,
waterborne
Mt. Assirik.
manner
However,
described
explanation
by
is
baboons,
and
yet
into
the
The
water.
irrelevant
show
(197,0),
to
levels
at
Mt. Assirik
in
duration
did
of
infestation
not
seem to
in
the
infection.
of
as have
behaviour
avoid
an
between
as long
for
adaptive
such
contact"
avoidance
schistosomiasis
cautious
drinks,
baboon
"minimal
the
unlearned
no
high
to
the
makes
which
the
when
in
at
towards
many parts
of
defaecating
or
water.
reflect
the
rivulets
and puddles
frequent
of
drinking
the
opportunities
in the rainy
from permanent water
between how thirsty
for
case,
despite
The difference
drink,
any
humans
Baboons
world.
urinating
In
been exposed
water
contaminated
the
Altmann
by baboons
drank
also
for
strategies
experienced
Mt. Assirik
at
and
therefore
Humans have probably
those
Altmann
with
different
evolved
from
parasites
baboons
ingested
and water
have
and
untenable.
is
parasite
lips
from,
pressures
baboons
different
had
sources
to
drink
may
from
season compared to the few they had to
sources,
the animals
at
in the dry.
were likely
correlation
to be and the length
which they drank was found by Nagel (1973).
6.1 6
A similar
of time
CHAPTER7: Feeding
INTRODUCTION
7.1: Previous
Work
have been no previous
There
of P. papio.
Reports
include
lists
food
selected
1976,
1976,
Hausfater
1978;
Westlund
tree
fruit,
of items, including
flowers,
shoots,
various
1978, Harding
Strum 1976).
any one site
P. hamadryas
1979) The feeding
Firstly,
does
it
live
studied.
sets
in
this
behaviour
for
a habitat
different
1974a, Hall
social
baboons, P. anubis,
baboons
of
to their
1963a, Hamilton
organisations
influenced
et al.
of T. gelada,
p. cynocephalus,
7.1
array,
in
or
greater
the baboons of Mt.
baboons of the genus Papio.
from other
in monkeys is significantly
(Clutton-Brock
that
three
the genus
to this
T. gelada,
Secondly,
from
for
baboons of
other
example
diet?
of
that
suggest
of interest
Does P. papio belong
monkeys,
1970, Hall
The diets
data
pattern
of
at
(Kummer 1968a, Stolba
of P. papio is therefore
feeding
foods
and animals
of plants
available
generalist
1978,
their
but
items,
of
1966a).
the
et al.
1977, Popp 1978, Post
1978, Rowell
These baboons may show adaptations
organisation
mushrooms, and
and Altmann
known, but
into
other
organs,
and storage
roots
1971, Altmann
on a more restricted-,
them apart
differing
al.
across the genus.
resemble
specialisation
Assirik
et
well
the
a wide variety
eating
buds, gum twigs,
and Maples
and
thorough
and
(eg Dunbar and Dunbar 1974c, Hamilton
1976, Post
ecology
is similar
Papio
leaves,
from up to about 100 species
not
Rhine
1978, ).
1975, Post
and leaf
flower
(Aldrich-Blake
are
1976),
Bearce
on
and Strum
and sedges;
Not only do they eat a wide variety
P. hamadryas, too, fits
reasons.
and
seeds, grasses,
1966, Hausfater
1963a,
tamarinds
baboons
studies
1975, Harding
baboons are omnivorous,
1976, Moreno-Black
gathered
are often
1975),
1973.19740
(Iwamoto
of animals
classes
1976, Jouventin
Hausfater
of diet
show that
Such studies
species
of
ecology
of
other
1975,1976;
Strum
exudates;
studies
quantitative
bark,
(eg meat; Harding
items
food
1962a, Harding
feeding
on the
published
behaviour
feeding
on the
(Hall
studies
previously
which
environment
Thirdly,
by feeding
social
behaviour
1978, Iwamoto 1978).
The
P. hamadryas, and the savanna
and P. Ursinus
are now well
documented,
of
P. papio
of
the
that
while
comparison
is
uncertain
still
feeding
P. papio
of
ecology
it
species
is most likely
it
organisation
by
and,
resembles,
most
in
T. gelada,
of
which, if
A
9).
chapter
those
with
indicate
P. hamadryas, and the savanna baboons will
other
(review
any, of these
implication,
social
which
havc.
to
7.2: Omnivory and Terrestriality
The proportion
low,
other
but
omnivorous,
Baboons
of
to eat
are
up in
trees
studied
which lives
them
(Etter
1973,
to
be
Jolly
(Crook
level
ground
at
and
1978).
1978, Post
opportunity
environment
populations
forage
to
with
from
many species
may therefore
serve to redress
diversity
the
of
genus
feed
1970,
Altmann
and
(Altmann
trees
as a forest
with
animal
trees,
few or scattered
1963a, Hamilton
baboons which
have
trees.
At
Assirik
of tree
(Baldwin
Papio,
this
balance
since
has been
Popp
is
therefore
a somewhat reduced
they
1979).
live
in
an
Data from Senegal
dietary
the
to
emphasize
and
baboons
to eat items from many species
7.2
Mt.
other
1978 (Kuiseb),
et al.
at
or in areas with
on baboon diet
literature
of
than [do]
baboons whose diet
However, most other
The current
towards
(Jouventin
ground
and P. hamadryas all
P. ursinus,
competent
(eg Hall
of trees
on the
1962a, Kummer 1968a, 1968b) and at one site
1965, Hall
1966a).
is rarely
in rainforest,
primarily
but also exploit
in open country
lived
opportunity
similarly
expect
life
principally
P. cynocephalus,
(Rowell
few species
biassed
behaves
P. papio
we should
exclusively
almost
"do not seem any less
species"
then,
1976 and Strum
Hausfater
terrestrial
and feeds
at ground level
DeVore and Hall
least
to
adapted
M.sphinx,
P. anubis,
largely
genus
feeds
Wrangham 1976).
1975).
Papio,
of
be
to
1968, Dunbar 1977a, Dunbar and Dunbar 1974c, Iwamoto 1975,
Aldrich-Blake
seen high
If
reviews).
tends
baboons
few animals.
well
T. gelada
1970).
the
for
diet
the
5% (see
than
less
for
and 161-163
baboons
matter
animal
accounting
normally
1976 pp 132-137
to
of
in
at
of trees.
this
site
have
the
7.2.1: Seasonality
of Diet
Since
feed
baboons
to
expected
sites
the
et al.
1975),
of a given
(Pan
troglodytes)
Hall
in
at
(P. anubis)
the same species
Mt. Assirik
albigena,
Colobus
badius,
prep,
Primates
with
types
food
Colobus
Colobus
1966a),
diet
foods
most
many of
ate
the baboons
at
primates
(eg
to other
and Dunbar
1974d;
Struhsaker
1967b,
aethiops,
changes
show seasonal
should
43
plant
populations
regarded
few
50% of
but
1978, Post
items
food
will
feeding
all
(eg Rowell
If
1964
few of
P. papio
make up the
type
food
on relatively
1978).
of
recorded
a single
as generalists
specialise
apparently
to them (Popp
example,
species,
(1978)
Oates
with
proportion
on a small
and 35% concerned
species,
are
all
diet
their
in
items
strongly
For
them.
on
relatively
eat
depend
to
Baboons
available
baboons,
do not
typically
a single
species.
but
other
resembles
bulk
their
of
at Mt. Assirik.
The predominance
baboons
at other
of
study
on fruit
in
their
diet
(data
from
1978,
Post
1978,
Harding
at Mt. Assirik,
in their
diet,
should
food,
seems to depend
sites
general,
Popp
of
one class
the more wooded or forested
site
If
Dunbar
guereza,
their
diets
feeding
concerned
that
1978,
Tanzania,
of Items in Diet
available
guereza
records
diet
and
and, indeed,
baboons,
prep),
varied
but
frequency,
equal
the
in
Importance
7.2.2: Relative
from
troglodytes)
to rainfall.
related
the
(Pan
Park,
1975; Cercopithecus
Struhsaker
Harrison
cyclicity
1979,
Chimpanzees
National
chimpanzees
of
(Baldwin
(Ransom 1971, Wrangham 1975).
1975;
Waser
foods
plant
to
be expected
not
would
some
(Chivers
asynchronously
show seasonal
to other
are similar
Most
rainfall.
of plants
Colobus
in
Assirik
Gombe Stream
at
and fruit
diet.
At
become available.
on such species
their
to
related
prep)
baboons
Mt.
items
leaf
species
in
variation
of plant
plant
feeding
and primates
show seasonal
Lee
as different
change
plants
variety
on a wide
may be
diet
their
items
the
Dunbar
include
as do the chimpanzees
site,
the
partly
the
more the
in
of a large
1979).
(Baldwin
7.3
6
the diet
baboons
the
of
In
type.
1974. Hamilton
P="
fruits
there
upon habitat
Dunbar
and
1976).
in
such as fruit,
depend
et
al.
well-wooded
number of
species
7.3: Comparison
Several
relied
to
items
found
studies
discarded
hypothesis
the
food
types
that
lib.,
ad
quantitative
study
is
there
diet
Feeding
between
comparison.
have
on occasion
is,
food
site,
for
are used to
of
proportions
and estimated
remains
for
used
not
were
remains
this
have
a feeding
Data from Mt. Assirik
by feeding
1979)
that
remains,
had left
animals
estimated
for
except
1975, Baldwin
no relationship
observation.
analysis
Remains"
on feeding
of a species.
in the monthly
direct,
by
Jouventin
extent
the
after
the diet
about
test
lesser
or
"Feeding
with
(eg
on primates
a greater
information
Observation"
of "Direct
7.4: Boulder-Rolling
Baboons
at
several
(Altmann
stones
three
search
insects
is
of
Fady (1972)
that
observed
of their
Two
was seen correlated
of
beneath
baboons
were
7.4.1: Competition
feeding
management
Secondly,
of
Grand
Pare
suggested
that
the
(1977)
explosion
and
causing
certainly
the
most
responsible
of
cockroach.
in
over
the
frequency
they
did
with
of finding
indicate
might
foraging
which
the
any category
it
was that
the
over
boulders
at
what
turn
under
to conceal
food
objects?
Impact
baboons
is
National
the
ecological
du
Niokolo-Koba.
are undergoing
damage
to
primate
for
damage
7.4
of
in
to
the
trees
the
to
the
Dupuy
and
importance
great
baboons
numerous
more
baboons
most likely
of a size
ecology
Verschuren
probably
for.
and Ecological
the
the probability
with
This
boulders
in
changes
boulder?
searching
random or choose
The
watching
were seasonal
the
up to
of
boulders
in the GPN de NK rolled
from
behaviour
object
rocks
flightless
of
that
reported
roll
in
stones
foraging.
arose
Firstly,
and two species
P. papio
daily
questions
boulders.
termites
seen to
were
of
(1962a)
Hall
(1959)
Bolwig
turning
the
while
Africa
seen
literature.
the
[P. ursinus]
baboons
South-West
47x31x1Ocm in search
the course
in
a common activity",
in
baboons
to turn
over
Wt
Lthere are only
comm),
been
comm, Strum pers
behaviour
"among wild
commented
P. ursinus
this
of
that
of
comm, Popp pers
pers
reports
sites
park.
park,
and
and
bushes
a demographic
Baboons
are
than
are
therefore
was any
other
first
feeding
for
evidence
study
the
to
extent
one of the rarest
both
foods
(Baldwin
area,
the
foods
and if
diet
broader
baboons
than
baboons
occasions
approached
when
by
displacing
baboons
considered
to
Gombe
feed
were
is
which
1979; this
for
directly
and if
on two
Collins
diet
at
comm).
pers
than
did
are
have a
be
to
true,
Mt. Assirik,
in
site
and
On both
occasions.
seemed
Baboons
overlap
Chimpanzees
a feeding
is
this
baboons
over
the same
should
sites
dominant.
Chimpanzees
Mt. Assirik,
1971;
(Baldwin
at feeding
same tree
the
and
the numbers of chimpanzees.
seen to move away from
to have a broader
expected
in
be dominant
(Ransom
one
chimpanzees.
at
competed
subordinate
the
should
seen
were
which
to
they
field
overlapped
which
baboons
of
the
over
widely
ranged
of
same species
that where two species whose diets
is
(1977).
Mt. Assirik
at
the
a home range
having
have limited
might
(P. papio)
many of
ate
of many troops
Morse (1974) suggested
sympatric,
(Pan
troglodytes),
chimpanzees
They both
study).
home ranges
the species
environment;
to by Dupuy and Verschuren
and both
apparently
as chimpanzees,
influence
and baboons
Baboons were numerous,
study).
their
damage referred
chimpanzees
encompassed
baboons
some
in the park, and whether or not they are likely
1979, this
the
baboons, and provides
which
frugivorous,
largely
This is
they eat.
out on these
(Pan troglodytes)
Chimpanzees
certainly
almost
are
whose fruits
they compete with
primates
to cause the ecological
were
carried
in how far
specifically,
hand they
other
for many of the plants
agents of dispersal
the
On the
non-human primate.
the
tree
capable
of
may reasonably
be
they
at
as
might
were
be
therefore
chimpanzees.
METHODS
7.5: Collection
of Data
Feeding
behaviour
included
observations"
whether
baboon
the
or
not
these
was seen eating
species
of
plant
was recorded
all
objects
sightings
were
I recorded
or animal,
by two methods.
the
of
time,
feeding
Where
identified.
and, where
7.5
baboons
the
"direct
Firstly,
on
objects
when
possible,
age and sex of the
relevant,
the
part
of
the
a
baboon,
plant
on which it
was feeding,
attempt
to make records
animals
feeding
of feeding
that
and its
feeding
of
food although
food were recorded.
I therefore
intervals.
or more during
Thus a single
of 2607 direct
this
observations
"feeding
chapter
observation
includes
although
baboons.
actively
taken
this
find
animal
effort
for
carefully
of
In the remainder
of
of the baboons,
diet"
of the diet
is
category
included:
found
in
litter
holes,
areas,
"boulder
or
rolling
as efforts
looking
restricted
These areas I call
leaf
digging
at, or apparently
of the
of baboons eating
observations
ground,
neither
The feeding
remains.
both of which were regarded
the
from direct
with
to
the
and moving
for,
food.
but
not
(Anecdote's
and
often
fields".
of the Data
of feeding,
the data from two troops
were combined.
The feeding
final
food items
stated,
composition
In the latter
tugging
were typically
In the analysis
Fable's)
to all
food.
observations
explicitly
to the "apparent
observing
pans.
that
collected
from feeding
the true
logs,
inspecting,
always on laterite
7.5.2: Analysis
food.
foods.
prey, making sweeping movements through
trees,
Boulders
where
refers
or rotten
hands while
through
except
may not represent
searching
boulders
to
to be equivalent
Foraging
over
and
1015
the data
refers
record"
of 1
was not established.
by
plant
on
The data consist
observation.
supplemented
to data collected
nor refers
is
record
were
baboons,
of
records
three Adansonia digitata
whose identity
on objects
eaten or chewed and discarded
partially
of baboons
of which 2024 concerned identified
observations,
eating
by an interval
separated
an hour, eating
was scored as a single
and 583 concerned feeding
Direct
animals
feeding
scored as separate
several
observation
which I had not seen any baboon eating
baboon, in view for
in succession
fruit
as a single
age and sex of all
of feeding
In the
one another,
of
I could not depend on being in sight
the same food any observations
minute
independent
the
to the food.
relative
at the same source were regarded
on that
at regular
behaviour
records
used in
12 months of the study.
7.6
this
analysis
were collected
over
the
the
commonly
found.
This
S. Hall
to occur
leafing,
the
against
flowering,
and fruiting
in the field
anthropoidea
were known
These
area.
in
the
foods
the
area.
major
of
of the availability
data give an indication
most
an independent
bushes and vines whose products
of the four
in the diets
were
species
(pers comm).
recorded
of trees,
food
various
was checked
assessment
and others
of 20 species
the
which
assessment by Harrison
subjective
cycles
in
habitats
determine
to
were
used
assessment
data and a subjective
Both the SAPP transect
7.5.3: Dimensions of Boulders
The dimensions
and
the
dimensions
which
were
the
by
recorded
The
mean of
geometric
whose sizes
these
included
the
longest
axis
to the first
orthogonal
axis
was taken
dimensions
three
Dimensions
recorded.
as
the
in the wet season and 50 in the dry season,
over 100 boulders
I turned
longest
the
through
fields
boulder
then
The
recorded.
were
also
axis,
by baboons,
over
(GMD) of the boulder.
mean dimension
geometric
were
longest
the
and then
them
at random from
been foraging,
measuring
to the first,
beneath
chosen
had recently
baboons
had been rolled
that
substratum
131 boulders,
of
orthogonal
two.
of
of
nature
72 boulders
the
selected
range
baboons.
by the
items of food found beneath these boulders
All
potential
were recorded.
7.6: Damage to Plants
Faecal
samples
undamaged their
methods
passage
7.7: Comparison
Dietary
agents
of Diets
data
samples (Baldwin
for
baboon gut.
survive
physically
My notes
on feeding
assessed
dispersal
baboons
the
of
agents
as
of
of seeds
damage done
to
the
plant.
of the plants.
of Baboons and Chimpanzees
chimpanzees
1979).
one food type, and could
longitudinal
the
ever
seeds
I
any
the importance
and as destructive
whether
through
included
always
subjectively
showed
at Mt. Assirik
Each faecal
therefore
sample of the diet
sample normally
be considered
of a single
7.7
faecal
based
upon
are
contained
to reflect
chimpanzee.
more than
a fractional
The proportion
of
food
major
types
the
considering
month,
and
number
of
by chimpanzees
eaten
total
the
number of samples
proportion
month by month (using
direct
for
over
the
1977.
baboons
year
with
which
feeding
month's
for
record
for
items
by
from
diet
the
were
the two species
in that
type
the
compared
same month of
of chimpanzees
in
highest
were also
exist
the
year's
The frequency
items
three
top
the
summing
were
compared.
among the
ranked
data
These
ranking
and chimpanzees
foods
calculated
data
the
with
food
food
class.
No data
The fifteen
of
food
by
be estimated
that
containing
classes
tau)
baboons.
diet
apparent
that
Kendall's
of
observation
October
major
of
containing
samples
in each month could
for
any
compared.
RESULTS
7.8: Omnivory
Baboons at Mt. Assirik
dependent on plant
1.2% eating
for
animals
Known
insects,
neonate
seen to
(Table 7.1).
direct
were largely
they
of feeding
observations
identifiable
Of all
5.4% were of baboons rolling
invertebrate
although
Of the 2607 direct
material.
77% were of baboons eating
or foraging,
foraging,
were omnivorous,
of feeding
observations
over boulders,
and
of plants
parts
or
searching
apparently
prey.
animal
a lizard,
be eaten
termites,
and two hares.
(perhaps
gazelle
included
prey
One other
Tragelaphus
during
the
course
the
of
was
rufus)
(McGrew et al.
study
a
possibly
vertebrate,
or Cephalophus
scriptus
other
and
grasshoppers,
1980)
(Table 7.2).
The number
wet
than
season
directed
at
feeding
record
in
24% of
apparently
observations
in
the
The proportion
p<0.001).
than
of
discovering
the
all
(chi
all
of
for
foraging
feeding
animals.
7.8
with
was also
= 74.16;
df
and
squared
effort
together
animals,
on
on animals
(chi
season
squared
observations
searching
dry
animals,
concerning
dry
of
feeding
higher
= 1;
was apparently
of
proportion
in
the
wet
p<0.001).
foraging
the
1;
=
df
= 18.28;
which
the
in
was higher
concerned
In
the
season
June,
baboons
Table 7.1:
in
Number of Observations
Categories
of Various
Jan
Ident
(%)
Feb
Mar
Apr
May Jun
Jul
Feeding Record
of Food
Aug
Sep
Oct
Nov
Dec Total
165 115 205 254 109 159 202 150 168 135 154 124
66.8 75.2 80.4 81.2 78.4 59.6 67.8 73.2 78.1 80.8 83.2 76.1
1940
74.4
2.5
54
2.1
0.6
30
1.2
28
34
45
42
28
72
30
51
27
104
50
Unknown 72
29.1 19.6 16.5 16.3 19.4 39.0 24.2 24.4 20.9 16.8 15.1 20.9
item
583
22.4
163
2607
888730630434
Part kn.
3.1 2.2
sp unk. 3.2 5.2
Animals
200104
0.8 0.0
247
Total
Observations
153
0.0
255
0.3
313
2.2
0.0
0.0
2.0
1.5
1.5
18
6.0
22001
1.0 0.9
267
139
298
205
0.0
215
2.4
0.0
167
1.6
0.0
185
Columns show monthly variation
in sample size.
to
baboons
Rows show various categories
food
were
seen
of
which
eat.
Ident = identified
food item
Part kn. sp unk. = identified
species
part from unidentified
Animals = all identified
animal food
Unknown item = neither
part nor species were identified
In each cell,
top figure
is sample size, lower figure represents
in that month.
percentage of observations
Table 7.2:
in Diet
Animals Known to be Included
of Baboons at Mt. Assirik
No. of direct
Observations
Species
possibly
or
Termites
Grasshoppers
other INSECTA
Lepus crawshayi
Agama agama
Tragelaphus
scriptus
Cephalophus
rufus
7.9
19
*
2
6
2
1
1
Figure
7.1:
Cumulative
items
number of food
diet
of baboons
against
plotted
feeding
known, to be included
number of observations
in
0
0)
s4 °'
I
Q4
A
Q)
Zb
0
o
,>w
c
00
U
1000
Sample Size
2000
the
of
F
Figure
7.2:
Percentage
of this
known to be eaten
'v
0
C)
x"
known food
month's
in previous
month
items
which
were
100
C
. r.,
Mar
Apr
'o
Jan
Feb
"ý+
50Q
0
Oct
Nov
Jul
Aug
Sep
Apr
May
Nov
Dec
Dec
Jan
o
U
Feb
Mal
Oct
Jun
Jul
Aug
Sep
May
Jun
Wet
Season
Dry
Season
Oct
Nov
also
Figure
Percent
'C
7.3:
of observations
of
feeding
on fruit
and
100
O
V
O
CD
'o
Q)
N
P4 50
0
+1
C)
G)
NDJF
Dry
Season
M'. AM
-JWet
JAS0
Season
seeds
7.8.2:
Breadth
Baboons
of Diet
58 species
than
species
of
(Table
7.9)
of
(including
plants
although
95 different
most
of
record.
increased
as sample
number
the
to
items
list
of
least
2,
known
little
to
throughout
known
of
items
food
plant
included
the
study
recorded
were
the
feeding
in
(Fig
diet
the
7.1).
in
once
roughly
more
many more,
to
be
foods
from
food
plant
and probably
contributed
increased
size
was added
them
food
of
classes
at
all,
In
grass.
The
new item
14 different
least
at
ate
One
25
every
observations.
In every
the
but
month,
previous
previous
seen to be eaten
Fifty
food types
that
they had eaten
food
that
they
45%±8.8%;
was
in
in
the
previous
and Lannea
month
seeds and fruits
Borassus
of
feeding
food
(Fig
7.2).
any two
the
had not
Oxytenanthera
of
Vitex
of
items
been
items
digitata,
of Adansonia
in
eaten
of
of
that
types
fruits
aethiopum;
half
in the
by
the nine
up
made
was
fruits
shoots
had not
record
roughly
to
record
(the
acida;
the
is,
each month belonged
most to the record
senegalensis,
in
that
per cent of the feeding
contributing
types
The mean overlap
months
be eaten
included
also
month.
consecutive
known to
baboons ate
month,
Saba
abyssinica;
and
madiensis
Spondias mombin, and seeds of Combretum spp. )
Seven of
the
items
nine
Sixty-six
most frequently
two,
seeds.
their
fruit,
fruit
and seeds was reflected
in
fruit
made up 60% of the
feeding
63.4% in
of the food species
per cent
26% were exploited
while
the dry),
while
total
for
the
17% of the
seeds (15.4% in the wet season,
7.8.3:
Seasonality
both
wet
squared
and
seeds
squared
feeding
record
were fruit,
and
for
were exploited
seeds.
The emphasis on
Baboons eating
record.
(57.2% in the wet season,
record
was made on baboons
18.9% in the dry)
(Fig
7.3).
of Diet
and seeds
contributed
and dry
seasons
(chi
= 3.43;
df
ns
together
= 5.87;
their
feeding
eating
Fruit
recorded
= 1;
almost
= 1;
= 3.09;
squared
for
was higher
df
equally
but
seeds),
in
p<0.02).
7.10
the
dry
to
df
the
season
A strong
the
feeding
= 1; ns for
contribution
than
seasonal
in
record
fruit;
of
the
in
wet
difference,
chi
fruit
(chi
but
in
the
baboons
were
These
different
Fig
in
month
dry;
classes
bamboo and
from
foods
The proportions
feeding
(Table
month
many of
of
were
apparently
Influence
of
record
as
to
Shoot and Stem
Flush and Leaves
Gum
Bulb
Buds and Flowers
Roots
Other
Fruit
Seeds
Bark
0.96
0.76
0.44
1.10
0.58
0.82
0.48
0.23
0.10
0.33
(Fruit
0.67
and Seeds
The proportion
shoots
of
that
while
together)
86%), the
summarised
included
types
in
rainfall
that
of
(Table
Direction
of correln.
fruit
in the feeding
fruit
was eaten,
of
< 0.002)
While
roots,
feeding
tau = -0.78;
the more seeds were eaten.
gum, bulbs,
(Kendall's
fruit
independent
0.001
0.001
0.05
0.001
0.005
0.001
0.02
0.15
0.33
0.07
leaves,
when rainfall
7.3).
7.11
3.0
rainfall
buds and flowers,
(Kendall's
<
<
<
<
<
<
<
flushing
with
seeds was not
record
4.3
3.5
2.0
2.2
2.6
3.7
2.2
1.1
0.5
1.5
-
and stems,
Significance
z
+
+
+
+
none
none
none
80%
to each month's
median
of
a
proportion
between
class in diet
correlation
of food
was greater
p always < 0.005)
contributed
in
on Diet
Rainfall
correlated
was positively
always < 0.05),
tau,
the
to
related
Kendall's
tau
Food Class
seeds
food
different
on
7.3).
Kendall's
tau for
rainfall
and proportion
(taken
is
This
month.
the
a dependence
of
a result
Table 7.3:
and animals
in
= 1; p<0.001).
df
came about
changes
(13.2%
stems
and
the
to which
degree
the
shoots
grass
= 126.9;
squared
of
apparent
was also
eat
chi
seasonal
7.6.
the
to
seen
0.4% in
wet;
direction,
opposite
in
tau,
and fruits
was lower
of
together
(range
the proportion
p<0.001);
and
(Kendall's
and seeds
record
P
66% of
the less
Since
fruits
some
Adansonia
digitata
proportions
of
items.
Thus
(1Ox7x3cm)
The
in
the
dry
baboons
of fruit
7.8.4: Distribution
high
food
field
species
although
any one food
type,
normally
with
woodland,
but
woodland
(Harrison
this
the
of fruit
in the dry
large
month.
seeds
of
to
greatly
fruit
all
species
eaten
allowed
the
season.
96% of
varied
be found
might
distributed
were not
(Table
type
habitat
with
evenly
one habitat
in more than
densities.
be found
could
both
in
be found
in
(61%)
the
to
Most
prep).
of
and open
closed
closed
than
in
known food
open
species
in
5
6%
found
or more
were
and about
in the number of food species
differed
squared
by baboons
61.83;
df = 1; p<0.001),
=
with
which might
71% of food
in closed woodland and only 2% on grassy plateaux.
gallery
grassland,
contained
intake
more likely
in
Habitats
Riverine
scrub
single
in more than one habitat,
occurring
contained
the
in
(36.1% of
season
to
to
food
single
contributed
and this
species
digitata
was far
be found in them (chi
species
digitata
distribution
different
Thus Adansonia
7.5).
due
wet),
eaten
Their
area.
7.4),
(Table
the
dry
to
germinated
in
variation
was due
largely
was
the
of Foods by Habitat
The plant
occurred
in the
eaten
their
some of
August
(eg
some months
largely
which
of Adansonia
1.3% of
season,
the
February
The fruit
7.9),
in
eating
aethiopum,
in
eating
to maintain
across
seed
Borassus
of
africana.
the
in
Table
in
can be attributed
classes
peak
seed
the proportion
in
in December,
certain
important
overwhelmingly
fruit
seeds
peak
Afzelia
were
all
forest
known
combretum
and open woodland
and closed
plant
and
food
grass
19.6%.
7.12
species,
on
plateaux,
in
contrast
which
between
to
between
them
scrub,
them
Table
7. u:
Proportion
of Food Species Growing
in One, Two, or More Habitats
% of Species
No of Habitats
in which sp.
was found
39.2
54.9
9.8
3.9
2.0
Table
7.5:
Distribution
The table
RGF
Plant
of
Food Species
shows the habitat
Woods
Acacia dudgeoni
Acacia sp
Adansonia
digitata
Afzelia
africana
Alophylus
africanus
in which various food species are most
Dry
For
Species
Food
Habitat
found
commonly
Grass
Comb
Scrub
Bush
**
Anona senegalensis
Bombax costatum
Borassus aethiopum
Cassia sieberiana
Ceiba pentandra
*
*
Cissus populnea
Cola cordifolia
Cola sp. unknown
Combretum tomentosum?
Cordia myxa
*
Cordyla
Diospyros
pinnata
mespiliformis
Erythrophloem
suaveolens
Ficus leucardii
Ficus umbellata
*
*
*
ft
**
*
*
**
*
*
*
*
Gardenia
erubescens
Grewia lasiodiscus
Hexalobus
monopetalus
Kaempferia
aethiopica
Khaya senegalensis
7.13
Table
7.5:
Distribution
Food Species
Landolphia
heudelotti
Lannea acida
Lannea microcarpa
Malacantha
alnifolia
Manilkara
multinervis
Dry For
of
Plant
RGF
Food Species
Habitat
Woods Scrub
a
Nauclea latifolia
Oncoba spinosa
Oxytenanthera
abyssinica
Parkia
biglobosa
Pericopsis
laxifolia
*
Piliostigma
thoninngii
Pseudospondias
microcarpa
Pterocarpus
erinaceous
Raphia sudanica
Saba senegalensis
*
*
*
*
***
*
Spondias mombin
Sterculia
setigera
Strychnos
spinosa
Tamarindus
indica
Treculia
africana
Trichilia
Urtica
prieuriana
sp. unknown
Vitex madiensis
Zizyphus mucronata?
sp. of Meliaceae family
sp. of
*
**
Sapotaceae family
grass species
(1)
sp. unknown
(2)
unknown
sp.
(3)
unknown
sp.
******
(4)
unknown
sp.
*
KEY:
Dry For
RGF
=
Woods =
Scrub =
Combr =
Grass =
*
may be with bamboo understory
= Dry Forest,
Gallery
Riverine
Forest
Open Woods
Scrub
Combretum Scrub on Plateau
Grass on Plateau
7.14
(cont)
Bush
Comb Grass
hI
Figure
7.4:
Monthly
variation
trees
and bushes
in
percent,
of
observations
of
feeding
100,
10
Trees
0
ü
C)
O
Bushes
10
a,
w 50
P
4-4
op
I
0
a
C)
V
o'
a)
o"
NDJFMAMJJAS
Dry
Season
Wet Season
0
in
Figure
Monthly
various
7.5:
variation
types
of
in percent
plants
of
observations
feeding
of
4010
0
U
A
i1.f%
C,
CAD
it
St
A
4)
it
YO 204-4
Ji
f'
"1")
JFMAM
ND
Dry
Season
J
JAS0
Wet season
ýf
in
7.8.5: Arboreality
Of all
preponderance
all
feeding
wet
degree
forbs
of trees
observations
food
(42.7%)
to which
differed
(chi
were of
the
squared
baboons
67% grew as trees
species,
in
was reflected
to be seen feeding
more likely
the
known plant
in
the
baboons
= 52.7;
were
seen
from month to month (Fig
the
record,
in
feeding
in
trees
feeding
dry
(Table
Baboons
(80.5%)
df
= 1; p<0.001)
(Fig
to
feed
grass,
in
bushes,
7.4 and 7.5).
Table 7.6:
Growth
Forms of
Food Species
BUSHES
Acacia dudgeons
Combretum
Cordia myxa
Gardenia
erubescens
Grewia lasiodiscus
Hexalobus
monopetalus
Meliaceae
sp
iuglandifolia
Sorindeia
Strychnos
spinosa
Trichilia
prieuriana
Vitex
madiensis
TREES
Acacia sp
digitata
Adansonia
Afzelia
africana
Allophylus
africanus
Bombax costatum
Borassus aethiopum
Cassia siberiana
Ceiba pentandra
Cola cordifolia
Cola sp.
Cordyla
pinnata
Diospyros
mespiliformis
Erythrophloem
suaveolens
Ficus leucardii
Ficus umbellata
Khaya senegalensis
Lannea acida
Lannea microcarpa
Malacantha
alnifolia
Manilkara
multinervis
Oncoba spinosa
biglobosa
Parkia
laxifolia
Pericopsis
Piliostigma
thonningii
Pseudospondias
microcarpa
Pterocarpus
erinaceous
Raphia sudanica
Spondias mombin
Sterculia
setigera
indica
Tamarindus
Treculia
africana
Zizyphus
mucronata?
Zizyphus
spina-christi
Sapotaceae
sp.
FORB LEVEL
Kaempferia aethiopica
Urtica sp.
GRASSES
Oxytenanthera
abyssinica
VINES
Landolphia heudelotii
Nauclea latifolia
Saba senegalensis
7.15
This
where 61.6% of
trees.
season
7.6).
were
than
7.4).
vines
in
The
or
The proportion
trees
declined
steadily
of
the wet season.
in
bushes
of
the
and its
the
the
in
Feeding
the
wet
Saba
when
in
(Kendall's
trees
tau
positively
tau
= 0.68; p<0.002
bamboo shoots
The
obscure.
correlated
wet
frequency
month's
frequency
correlated
with
the
7.8.6: Direct
direct
feeding
observation
relative
order
next
and
importance
the
same month,
tau
= 0.68; p<0.002).
were
to
seen
(Kendall's
rain
month's
feed
Kendall's
and
rainfall
is
forbs
with
feeding
seen
were
and growth
appearance
were
rainfall
and
between
correlation
baboons
month's
but
more
bushes
in
well
the
with
On the other
in
tau;
vines
of
hand,
was closely
These
p<0.001).
later.
of
and Feeding Remains
the
20 food
feeding
observation
remains.
they
the
in
rainfall
which
Observation
The rank
but
which
(Kendall's
rain
with
season,
with
with
are discussed
results
the
the
poorly
previous
the
in
in
baboons were seen
p<0.002
by the
influenced
strongly
feeding
bamboo and forbs
The correlation
respectively).
probably
= 0.71;
tau
in
were abundant.
that
for
those
while
while
which
with
early
record
fruit,
in
record
fruits
record.
when shoots
correlated
(Kendall's
correlated
bamboo is
feeding
type,
madiensis
feeding
in
was
food
this
Vitex
the
in the wet season,
p<0.01),
-0.63;
37% of
up to
Vitex
species,
concerned
few ripe
due to
entirely
a single
record
start
in October
reached
0.6% to the feeding
only
was negatively
=
of
the
with
was almost
fruit
month
feeding
spent
sharply
in bushes
November
senegalensis
the
of
of time
and fell
last
the
until
proportion
very
were
baboons were using
which
season
feeding
on the
occupied
bamboo was most common late
feeding
the
in
decline
contributed
The proportion
dry
34% of the feeding
vines
season,
the
peak of
when there
plant
in
wet season
baboons
the
of
November,
remaining,
the
in
record
same time,
rapid
In October,
in
early
The high
subsequent
dependence
while
from
At the
season.
madiensis.
feeding
throughout
rose
dry
the
of
(Kendall's
feeding
of
remains
specific
types
record
tau
do
food
of
most
correlated
was not
= 0.63;
not
items
7.16
n=
give
in
frequent
20;
similar
the
diet.
occurrence
in
that
for
with
p=0.5).
estimates
However,
Direct
of
the
the
two
7.6:
Figure
Comparison
variation
of the monthly
in the feeding
derived
fruit
records
remains
and from feeding
;0
0
in the-percentage
of
from direct
observation
100
v
C
.,ý
41
w
50
w
0
+J
C)
u
Dry
Season
Wet Season
measures give similar
(Kendall's
monthly diet
of the percentage
estimates
tau = 0.62; n=
7.8.7: Comparison with other
species
data
from
strictly
in
comparable
the
table
therefore
Table
(Table
are
with
drawn
other
from
7.7).
Africa
South-West
and is not based on the diets
Africa,
(Fig 7.6).
12; p<0.001)
were seen to eat a similar
elsewhere
in
populations
in the
present
species
The baboons at Mt. Assirik
as were baboons
of fruit
in
of baboons
Cape Province,
in a single
Furthermore,
studies.
repeated
studies
area,
South
it
is not
of the
several
the
at
includes
list
Hall's
Since
and
number of plant
same site,
data
and are
independent.
not
7.7:
Number of Species Known to be Eaten
Theropithecus
by Papio,
# of
plant
spp.
and Mandrillus
at
field
various
sites
rank
# of
anim.
spp.
94
68
66
58
1
2
3
4
14+
M. sphinx
53
5
Jouventin-1975
P. anubis
42+
6
Dunbar and Dunbar 1974c
P. cynocephalus
25+
7
P. anubis
P. anubis
25+
24+
7
9
P. anubis
21+
10
P. cynocephalus
P. cynocephalus
T. gelada
20+
17+
15+
11
12
13
Baboon
species
P.
P.
P.
P.
ursinus
anubis
ursinus
papio
The true
number of
higher
undoubtedly
Hall-1963a
Popp 1978
Hall 1960
This Study
6+
Post 1978
13+
Harding 1976
Rowell 1966a
3+
species
(see Fig
Source of data
Aldrich-Blake
umbellata,
digitata,
Lannea
by baboons in this
eaten
7.1).
Few of
Pseudospondias
Annona senegalensis,
acids,
1971
Moreno-Black and Maples 1977
Altmann and Altmann 1970
Dunbar and Dunbar 1974c
the
species
baboons are known to be eaten by baboons elsewhere.
Adansonia
et al.
Malacantha
Diospyros
alnifolia,
microcarpa.
7.17
study
eaten
Exceptions
mespiliformis,
Nauclea
latifolia,
site
is
by these
include
Ficus
and
P. papio
Papio
in
its
P. ursinus
latter
This
dependence
heavy
and
from
baboon
despite
troops,
points
with
more
heavily
being
is
out,
"desert"
of
the
genus
two troops
7.8),
depending
as Kummer (1968a)
baboons
other
(Table
on trees
P. hamadryas
of
one
troop,
many savanna
different
greatly
was not
on
trees.
than
more arboreal
baboons.
7.8:
Table
Dependence
% of
Baboon
Species
of
Baboons
on Trees
Rank
from Trees
Diet
Food
for
Source
Data
of
P. ursinus
80.9
1
Hamilton
et al.
1976
P. ursinus
P. hamadryas
76.1
69.5
2
3
Hamilton et al.
Kummer 1968a
1976
P. papio
P. anubis
P. anubis
61.6
53.6
4
5
This study
Dunbar and Dunbar
39.4
32.8
10.0
4.7
0.0
6
7
8
9
10
P.
P.
P.
T.
cynocephalus
ursinus
anubis
gelada
Where there
more on trees
as there
in
the dry
matched pairs;
p<0.05)
more on trees
than
Canyon,
troops
rainfall
did
(Table
P. papio
in
7.8).
18mm; Awash, estimated
length
they
of
depend upon trees
the
dry
season
in their
(Kendall's
(Data from Dunbar and Sharman in prep).
7.18
(Wilcoxon
the wet
Two of the troops
in
were living
for
Papio
the
baboons
genus
of
of
to which
in the amount of food
baboons tended
was at Mt. Assirik,
season than
1974c
Popp 1978
Post 1978
Anderson 1980b
Dunbar and Dunbar 1974c
Dunbar and Dunbar 1975
difference
was a marked seasonal
from trees,
taken
the
of
rainfall
arid
annual
signed-rank
which depended
areas
(the
665mm), and,
which there
is
diet
tau = 0.62;
to depend
data,
for
eight
the degree
is correlated
n=8;
Kuiseb
with
p<0.05)
Table 7. 9:
Feeding Record of Baboons at Mt. Assirik
Full
names of species
are given
F.ju. cs rerres.t. I Pet C*&%tof ~M44
Species
Nov
Dec
Jan
Feb
Aca dud Base
Seed
Ada dig
Bud
Afz
All
afr
afr
f .. 4i y
1.3
0.8
3.6
1.7
Mar
Apr
May
1.0
0.8
0.9
1.5
0.4
0.9
2.0
0.4
6.1 10.4
2.4
1.2
0.4
1.6
Seed
Flsh
Bark
1.3
0.6
0.7
3.0
1.9
1.3
1.6
3.5
3.9 13.9 20.2
1.0
pop Frt
Leaf
1.3
Flwr
Seed
0.6
5.7
4.7
2.5 16.8 27.3
5.4
2.2
1.0
0.5
0.7
3.6
1.5
4.4 16.3
1.3
1.2
3.0
1.0
2.5
1.9
1.6
0.9
1.0
1.3
2.4
2.4
1.2
2.6
3.9
1.2
4.3
1.8
5.0
3.4
2.4
2.8
1.3
0.5
1.3
3.5
4.0
Dio mes Frt
Fie umb Frt
1.6
1.7
Cor pin Frt
Fie lee Frt
1.0
1.2
0.8
1.6
0.9
Cor myx Frt
sua Seed
8.0
O. u
0.6
Comb s
1.0
1.8
0.6
1.3
pen Bud
Flwr
Flsh
Seed
Oct
0.6
Seed
s
Ery
Sep
1.0
Col cor Frt
Cola
Aug
Frt
Bor aet Frt
Seed
Cis
Jul
1.5
Flwr
51.3 57.3 38.8 12.2
Frt
Bom cos Flwr
Seed
Cei
Jun
1.3
Ano sen Bud
Frt
Cas sie
ratae t.
Month
Part
Aca sp
in Table 7.5
3.9
5.6
8.5
7.0
0.5
0.4
3.2
1.8
0.9
1.0
1.2
0.9
2.4
7.3
7.8
1.0
3.9
2.8
1.8
3.5
1.5
2.8
3.7
3.3
0.7
0.5
1.3
2.5
2.4
2.7
1.5
Table
Species
7.9:
Feeding
Record
of
at
Mt.
Assirik
Month
Part
Nov
Dec
Gar eru Frt
Gre las Frt
Baboons
2.6
Jan
Feb
0.6
0.9
Mar
Apr
May
Jun
Jul
1.0
1.6
1.3
Kae aet Bulb
1.2
3.7
1.3
12.5
3.7
2.0
2.0
1.2
1.0
Gum
Bark
Lan heu Frt
6.1 30.7 17.3 13.8
Lan aci Frt
1.5
Lan mit Frt
8.3
0.9
1.3
1.5
3.0
0.7
0.7
1.3
5.9
0.7
1.3
0.6
1.2
Mai aln Frt
1.3
Man mul Frt
1.3
5.8
1.8
8.9
3.6
3.7
1.3
One s pi Flwr
Frt
0.7
1.2
Kha sen Flsh
Twig
Frt
Oct
0.7
Hex mon Frt
Nau lat
Sep
Aug
1.9
3.4
5.1
5.5
1.8 3.1
Oxy aby Flsh
Stem 1.3
1.0
Base
Root
0.6
Litt
2.4
8.7
Par big Flwr
Par big Bud
Flwr
Seed
Bark
Unkn
1.0
0.4
6.8
3.9
1.5
0.8
1.5
0.5
2.7
1.8
2
1.3
1.8 3.7
0.4
0.8
Shoot
1.5
0.8
2.8
0.4
9.2
7.5
8.4 14.7 18.5
0.9
0.9
0.9
1.8
0.9
0.9
0.6
2.2
Per lax Seed
P11 tho
Seed
5.2
0.6
5.5
3.5 10.7
6.?
7.8
2.0
Pse mic Frt
Pte eri Flwr
Seed
Leaf
Bud
13.9 10.7
2.0
0.9
0.6
4.0
0.8
1.9
1.0
2.7
0.6
Table 7.9:
Species
Feeding Record of Baboons at Mt. Assirik
Month
Part
Nov
Dec
Jan
Feb
Mar
Apr
May
Jun
Jul
Sab sen Frt
Seed
Flsh
3.9
12.2
25.7
0.5
1.6
0.9
Sor iua Frt
.
0.5
1.2
0.9
32.1
10.9
4.0
Soo mom Frt
Str
s pi Frt.
Tam ind
Seed
Tre
afr
Frt
pri
Frt
Tri
1.0
Frt
set
2.6
4.8
sp
Vit
mad Frt
Seed
3.1
0.6
0.9
0.9
1.3
3.2
2.4
2.6
Stem
Unkn
0.6
Sp Un 1 Frt
0.7
0.5
0.5
3.3
0.7
3.2
0.8
3.6
3.5
1.5
1.9
3.8
2.0
1.0
2.0
5.0
1.0
2.0
2.0
1.3
1.8
4.8
2.4
2.4
1.5
5.2
0.7
1.2
0.4
1.5
1.2
3.2
Sp Un 2 Frt
1.3
Sp Un 3 Base
0.7
Sp Un 4 Twig
0.6
Gum
1.9
Unkn 7.8
Unk Ori Leaf
Bark
4.8 34.1
1.5
1.3
1.3
5.8
Frt
1.0
1.0
Sho gra Frt
Leaf
Mel sp
2.2
2.0
0.6
0.5
Stem
Bud
Frt
0.7
0.7
Ele gra Leaf
Sap sp
8.9
0.6
Bark
Ziz spi Frt
1.3 25.6
2.7
4.0
0.5
1.2
Base
Urt
Oct
0.6
Rap sud Frt
Ste
Sep
Aug
0.7
0.8
0.6
0.9
1.4
1.0
1.3
0.7
Feeding Record of Baboons at Mt. Assirik
Table 7.9:
Part
Species
Month
Gum
Root
Jan
Dee
Nov
1.9
3.1
Feb
Mar
Apr
2.9
5.7
0.9
2.8
0.8
1.9
1.2
0.8
Ele Fae Seed
May
Aug
Jul
Jun
Sep
Oct
0.5
2.9
1.8
ANIMAL
0.8
Termites
Grasshoppers
Other Insect
Hare
1.1
2.5
0.6
1.3
4.0
0.9
2.7
0.4
0.4
0.6
Lizard
Observations
157
129
175
123
213
Roll bouldrs
1.6
4.3
3.6
2.6
1.6
226
155
170
139
3.6 21.3
6.6
5.5
7.0
3.0
0.4
logs
Roll
163
112
262
1.1
1.0
3.2
7.6
7.4 11.3
4.9 10.1
5.9
8.5
6.3 11.2
6.7 2.9
7.2
5.0
9.4
4.5
9.4 10.6
1.0 2.5
8.4
4.7
7.2
4.2
Digging
Salt lick
2.7
4.3
2.6
2.0
2.2
2.9
0.7
1.9
0.7
3.5
0.3
2.5
0.9
2.4
Observations
185 163 247
313
139 267
Other ground
Off ground
Boulder-Rolling:
7.9:
Laterite
(Chapter
forms
resting
Laterite
has no fracture
irregularly
cavities
The
in
small
turned
by standing
levering
step
largest
around
the
the
boulders
over
167
boulder,
up towards
upright
boulder
baboon
one
seizing
the
and
straddled
7.22
its
chest.
push
the
it
weathers
and when
provide
Baboons
collect.
Larger
far
The
boulder
it
rough,
indentations
over.
area
plate-
detritus
hand.
field
occasionally
are
and
plant
with
boulder
the
though
protruberances
over
the
and when it
boulders
or
the
of
many parts
plane,
the
can hide
boulders
in
spheroidal,
of
the
surface
animals
could
often
surfaces
which
turn
and
shaped,
a hard
298 205 215
Results
were abundant
2).
on
153 255
boulders
boulders.
like
3.2
0.4
boulders
edge with
baboon
were
one hand,
could
then
To roll
over
the
its
legs
and
with
Figure
Percent
7.7:
of monthly
feeding
record
concerning
boulder-rolling
0
E
.o
Ot
0
0)
120)
0_
NbJFMAM
Dry
Season
JAS0
-J
Wet
Season
vigorously
pulled
to pull
continued
Boulders
failed
its
with
while
hands until
back to lever
stepping
were occasionally
the boulder
abandoned after
to lift.
started
the boulder
up and over.
tugs
two strong
one or
It
had
to move them.
Boulder-rolling
feeding
record
was seen
in
record
frequently
in
69.6;
=
df = 1;
turned
over
by
boulder
(n=42).
two out
the
(Fig
wet
a boulder
and
suggesting
In
placed
data
boulder
the
for
the
in
dry
the
wet
one or
in
something
for
rolled
the
dry
its
season
had been rolled
over
mouth
are
(chi
not
occasions
Twentymaterial,
plant
(Table
previously
had
covered
available.
on crushed
more
that
previously
on 32% of
no
squared
a baboon
items
the
had
boulders
season
season,
more
(April)
month
Baboons
21% of
formed
and
one
(31%) had been resting
72 boulders
that
than
season
reached
Only
7.7).
p<0.001).
Comparable
of
(June).
month
boulder-rolling
of
the
one
157 occasions,
on
7.10).
Table 7.10:
Substratum
had been Resting
on which Boulders
Embedded in
ground
Boulders of similar
size
Crushed plant material
Gravel
Laterite
or large boulder
In
any single
boulders,
passage
boulder-field,
hundred
several
through
the
The distribution
might
selected
df = 5;
5
0
0
16
8
18
0
6
4
15
of
sometimes
consisting
be turned
by the
thousand
several
baboons
in
a single
field.
of the geometric
mean dimension
by baboons and sampled from boulder-fields
= 58.3;
Dry Season
Wet Season
Substratum
p<0.001)
few of the smallest
(Table
were dissimilar
7.11),
or largest
7.23
of boulders
largely
boulders
because
available
rolled
(chi
squared
the
baboons
to them.
Table 7.11:
Distribution
Geometric
of Sizes
<= 10
Mean Dimension
The nature
the
between
boulders
of
over
small
than
one animal
pupae
diameter,
in
cavities
(Table
item
the
dry
eggs
season
26%,
differed
6% of
the
of
webs
silk-like
pasted
spiderlings,
Of these
38
some form
26% covered
or
>= 30
53
12
boulders
densely-woven,
and strong,
boulder.
17
11
beneath
although
containing
the
In
26-30
21-25
24
18
I found
7.12).
animals,
16-20
13
22
which
(Table
mobile
including
1cm in
about
objects
seasons
concealed
life,
animal
of the
two
11-15
10
30
Rolled
by baboons
Available
in area
of Boulders
31% concealed
flat
more
7.12).
7.12:
Table
Possible
Numbers represent
Food Items
Boulders
Found Beneath
(one - zero sampling)
per cent of boulders
Spiders
Beetles
Araneae
Coleoptera
20
18
Ants
Formicidae
14
Termites
Isoptera
12
Snail
Cricket
Gastropoda
Orthoptera
10
8
Scorpion
Solifugid
Other invert.
Scorpiones
Solifugidae
Chilopoda,
5
Frog
Salentia
Diplopoda
In
(difference
squared
the
1
wet
between
25.1;
=
df
17
not scored
not scored
not scored
not
or very
season,
wet
2
1
8
etc.
*
Unidentified
Plant detritus
Seed
Web
Chrysalis
* very small
2
2
4
96
42
20
scored
8
fast
64% of
and dry
= 1; p<0.001),
the
boulders
seasons
statistically
and of
7.24
concealed
these
mobile
significant;
animals
chi
64%, 56% had more than
0
Figure
7.8:
Number
of
animals
found
beneath
L"oulders
of
different
sizes
6
6
ý.
4
0
.0
,2
0
10
20
Geometric
30 "
40
Mean Dimension
(cms)
50
60
7.9:
Figure
Relationship
found
debris
Many
Common
Few
o
between
beneath
and amount
of boulder
size
in the dry season
it
00
0
o
00
coo
0
00
00
000
00
0000
00
0
0 00
00
00
10
20.30
Increasing
7.10:
Figure
Relationship
found beneath
between
in
it
Size
40
O
Common
50
(Rank)
of'Boulder
of boulder
size
the dry season
Many
and
number'of
0"
O0
seeds
00
00
0000000
000O00
Trace
None
0
OOO
None
Few
plant
00 0000000
'O
Trace
00
of
O 00
0 00
00
000
10
Increasing
0000
0 000
00 00
0000
30,40
20
Size
of
Boulder
00
50
(Rank)
Figure
7.11:
Relationship
between
size
of
beneath
it
animal
bars are 95% binomial
Vertical
boulder
and
chance
confidence
of
limits
1
a)
CZ
c
N,
4 c3
C) ü)
M
r-i r,
Z c.
0
ca c
4-4
0V
0
a)
a
5
Geometric
25
35
15
Class Mark of
Mean Diameter
M
ýý
of
Boulder
40+
(ems)
finding
beneath them (Fig
one animal
Plant
from
and were
of
the
found
were
as occurring
boulders.
Much of
leaves,
small
or of
7.9.1:
Size
white,
the
dry
such
bark,
and the quantity
beneath
it
in
be "common"
to
plant
12%
under
was probably
included
straw,
In
wet
so on.
crushed,
season,
detritus
remains
and
dry
the
the
dried
season,
and mildewed
yellowing,
stems.
and Number of
boulder
the
of
of mats of
etiolated
there
season
(Figs
dried
of
Boulder
of
rest
boulders
enough
season,
consisted
often
grass,
the
Seeds, often
boulders.
all
the
frequently
dry
the
portions
matter
In
in
food;
as
nearly
42% of
under
scored
useless
plant
was found under
detritus
grasses,
7.8).
Objects
was no
the
size
of
or the number of seeds found
detritus
of plant
between
relationship
and 7.10).
7.9
In the wet season boulders
(chi
ones
beneath them than did larger
10cm GMD had fewer animals
than
smaller
6.54;
=
squared
n=1;
p<0.02)
(Fig 7.11).
Damage to Plants
7.10:
Much of
branches,
but
baboons
from
species,
and
acted
(fruits
Hanno undulata)
are
proportion
the
(Kendall's
of
tau;
foods
vegetable
not
diets
for
and
known
24.
They
5 species.
for
of Baboons and Chimpanzees
of Garcinia
fruit,
apparent
agents of dispersal
57
the
dispersal
of
fruiting
and seeds,
damaged 22 of
agents
as possible
56 different
three
Assirik,
flowers,
buds,
Baboons
as possible
breaking
involved
plants
destroyed
trees.
Comparison of Diets
Of the
in
also
living
both damaged and acted
7.11:
damage to
conspicuous
the
bark
stripped
food
the
ovalifolia,
known to
seeds,
of
be eaten
flowers,
baboons
p never < 0.05)
eaten
shoots,
by chimpanzees
Icacina
by the
bark,
and chimpanzees
(Table
L IS
7.13)
across
at
and
senegalensis,
The
baboons there.
and flushing
are
not
months;
Mt.
leaves
correlated
that
is,
the
7.12:
Figure
Comparison
be included
increasing
(Method
of
so that
no
known to
of cumulative
number of food items
in diet
and chimpanzees
of baboons
with
sample size. '
in the two species,
different
collecting
samples
is given
axis)
scale
on the horizontal
0
C)
a
H
00
0
0
w
10
b-600"s
4
4-4
0
3
.n
2
pa"424C
a)
H
U
k
Cumulative
Sample
Size
two
species
different
place
on different
emphasis
food
in
classes
each
month.
Table 7.13:
Proportions
of Major Food Classes in Diet of Chimpanzee
Correlated
with those in Diet of Baboon
Class
Kendall's
Food
of
Fruit
-0.37
Seeds
Flowers
Shoots
Bark
Flushing
In
each
the
leaves
-0.28
that
except
to
tend
correlations
recorded
among the three
The rank
order
baboons (Kendall's
only
one list
was ranked
for
8,
eaten
of
the
is
plotted
sampling.
eaten
by baboons
wider
sign
than
of
that
at
reaching
of
ns
is
no correlation,
various
species
there
that
items
of
in
Fig
will
be recorded
food
items
at
100
chapter),
The diet
Mt. Assirik.
7.2
and
of
known
7.12.
few new items
an asymptote.
for
appeared
(Baldwin
items
food
size
(this
any month.
in
list).
The number
over
were
not the same as that
is
sample
same site
chimpanzees
for
in the diet
35 plant
eat
contrast,
the
0.26
(Any item
16th in the other
that
By
ns
ns
ns
which
p=0.39).
increasing
against
0.75
0.10
0.18
which
with
Mt. Assirik.
at
indicates
curve
faecal
little
4)
Table
0.11 ns
chimpanzees
Chimpanzees were known to
Chapter
for
frequency
= 0.04;
tau
11
most common species
the list
of
significance
be negative.
shows the
7.14
bark,
of
n
11
11
11
11
11
-0.07
-0.48
-0.35
0.00
case,
Table
tau
are
the
baboons
1979,
to
be
The shape
by
further
known
to
curve
is
be
shows
probably
Table
7.14:
Frequenc
Items mos
(max.
with which Food Species is one of the three
in Monthly Feeding Record.
commonly recorded
freq.
Pan troglodytes
= 11; for P. papio = 12)
Pan troglodytes
Papio papio
Freq
Rank
Rank
Freq
for
Borassus aethiopum
Adansonia digitata
4
3
Saba senegalensis
Oxytenanthera
abyssinica
Lannea acida
3
1
1
2=
4
9=
4
3
3
2=
4_
4=
9=
2
2
6=
6_
6
1
1
8=
2
2
5=
5=
1
1
8=
8=
2
5=
1
1
1
1
8=
8=
8=
8=
2
5=
9=
9=
9=
9=
9=
-
-
Diospyros mespiliformis
Pterocarpus erinaceous
Ficus
5
4
1
-
spp.
Cola cordifolia
Hexalobus monopetalus
Piliostigma
thonningi
Afzelia
africana
Strychnos
spinosa
Sp. unknown
-
Leaves
Spondias mombin
Landolphia heudelotti
Grewia lasiodiscus
Tamarindus indica
Insects
Honey
5
2
1
1
1
1
1
DISCUSSION
7.12:
Biasses
In
in
a study
the
this
of
tended
seen eating
Feeding
drawn
to
foods
Erythrophloem
suaveolens
(eg
digitata
will
such
Adansonia
tend
as
to
is
and juveniles).
whose
fruit
seeds
or
by a moving
or
Borassus
the
blades,
baboon
africana
feeding
can
and are
7.27
of
be
seeds).
record.
collected
presumably
the
was
seeds)
that
or less
males)
processing
aethiopum
and Afzelia
in
(adult
Animals
The attention
collection
and
inevitable.
conspicuous
be overrepresented
grass
inconspicuously
bias
sort,
to be either
than most (adolescents
was
Record
or
were
timid
observer
noisy
(eg
conspicuous
These
Other
and
underestimated
foods
foods,
eaten
in
the feeding
based
the
on feeding
remains.
The degree
to which
feeding
it
since
in
Similar
record.
will
of
be seen well
This
details
of
ecology
for
it
the
different
biassed
the
in
food
rivals
in
results
could
item.
its
conclusions
major
on
of the feeding
biasses
underlying
months,
level
ground
at
a broad outline
gives
which
with
foraging
depend for
does not
Instead,
P. papio,
probably
and
across
constant
attention
identify
to
enough
the diet.
of
trees,
record
or under-represented
remain
observer's
in
however,
study,
over-
general
to the feeding
apply
is
conditions
baboons
seeing
not
often
the
Observation
item
in
not
for
compete
each month.
favour
a food
will
record
considerations
will
be
buds,
flowers,
less
important.
7.13:
and Nature
Breadth
The
baboons
seeds,
fruits,
stems,
bulbs,
fed.
they
Mt. Assirik
at
leaves,
flushing
roots,
They ate
of
diet,
more than
50% of
seasonally,
(62%)
diet
in
probably
by baboons,
that
baboons of
find
a significant
of this
for
leaf
stems,
twigs,
plants,
three
of
orders.
record
their
food
was apparently
to
the
genus Papio,
top
derived
but
10
from
changed
rainfall.
They also
while
of their
radiation
diverse
their
year
at
ate
from the
was derived
homogenous through
on which
and also
Despite
thus expand the range of plant
the adaptive
gum, base of
58 species
response
proportion
bark,
from the plants
as expected.
the
and
feeding
of
was not
These observations
eaten
the
flower
unknown, items
from
animals
Most
Their
trees.
shoots,
least
at
6 species
items.
ate
and other,
least
food
of Diet
support
adapted
foods known to be
previous
to a terrestrial
food in trees.
life,
The implications
of baboons are discussed
7. Z8
evidence
below.
7.13.1:
Omnivory and Troop Size
Uneven distribution
patchy
distribution
times
when certain
high-ranking,
This
isolated
stands
therefore
food sources
of Ficus
be forced
P. papio
of
occasions
processing
seeds)
that
defenses
of
unknown,
Erythrophloem
This
them from
P. papio
the
other
dry
when
its
at
in
availability
Baboons
at
(1975)
Strum
documented
during
in
other
showed
abandoned
competition
the
that
a habit
are
increase
of
potential
a plague
Mt. Assirik
seasonal
at
season.
This
insects
included
presumably
in
the
in
(pers
the
of
aethiopum
chemical
plant
Cola
sp.
Gardenia
on
pressure
within
the
troop
3),
while
(Chapter
in
In
primates.
other
was
high
large
troops.
or
insectivores;
Animals
carnivores
baboons
living
mammals,
insect
obs),
capable
may have allowed
factors
food
for
record
7.19
the
Kenya
Insect
the
peak
et
change
between
the
in
the
wet
(1978)
al.
in
Botswana
is
abundance
occurring
and
acquired
by P. ursinus
prey.
with
in
and Hamilton
insect-eating
accounts
feeding
did
facultative
hunting
a major
than
any
a major
dudgeoni,
them to live
of
P. anubis
of
these
near
competitive
fragmented
allowed
Consumption
sites
of
if
aethiopica,
reduce
over
troops
wet season
Seasonality
7.13.2:
All
densities
highest,
the
to
few
on
or
would
The wide
Borassus
Acacia
Kaempferia
served
species.
higher
at
season,
presumably
then
may have
(eg
with
animals
also
mechanical
(eg
foods
particular
that
were
not
overcome
suaveolens,
primate
live
to
to
able
potential
other
were
primates
may be
spread of the
animals
area
a
alternative
sources.
meant
field
this
other
possibly
and were
erubescens).
in
in
low-ranking
for
when
and
food
may have
available
there
Subordinate
alternative
Assirik
Baboons
foods
for
with
by a few, presumably
Mt. Assirik,
at
that
the current
within
spp and Lannea spp.
Mt.
food
site.
occur
to search
at
was no
feeding
to
1978)
al.
food are not available
was seen
implies
can be monopolised
et
together
habitats
across
themselves
(Hamilton
animals,
troop.
diet
of the habitats
of that
sources
of food species
in
highly
the
proportions
and dry
wet
of
seasons.
That the four
instances
be coincidental,
might
accidentally
prey
the
frequency
vertebrates
of eating
or,
since
baboons normally
(Harding
1973,
Hausfater
baboons
might
with
which
in the wet season
occurred
discover
1976,
1976),
Strum
stumble
on
vertebrate
reflect
in
prey
the
wet
season.
7.13.3:
Seasonal
and Annual
The influence
the
in
of
predictable,
times
of
temporary
may have permitted
timing
observation
that
was correlated
as
A similar
Saba
the
the
with
dependence
in
the
Hexalobus
crops
single
fruit
Pseudospondias
previous
during
and
years
subsequent
and
overwhelmingly
patterns
and
(Tutin,
Harrison
fail
species
(pers
failed
entirely
subsequent
this
years
food
social
pers
comm).
for
study
the
organisation
7.30
to
fruit
baboons,
was
fruit,
was
influencing
during
species,
its
strong
if
the
year.
any
out,
carried
poor
very
produced
with
feeding
of
in
Harrison
Baldwin,
madiensis.
variation
fruit
or
compared
Tutin,
obs,
flower,
food
such
inter-annual
erinaceous
respectively,
perhaps
in
feeding
Pterocarpus
microcarpa
important
leaf,
this
which
and
seeds
to
Vitex
Conversely,
responsible.
Thus
for
a single
again,
rainfall;
an
interpreted
frequency
of
correlation
may result
types.
food
bush,
of
at
had
also
was better
rainfall
items
on bushes
baboons
the
turn
Temporary
species
of
to
in
diet.
on the
species
month's
species
year
the
largely
was
monopetalus
of
for
subsequent
preferred
certain
Thus
on
month's
on a single
holds
senegalensis,
by
This
or a few,
on one,
classes
leading
cycles,
1968).
a single
entire
Predictability
area.
plant
from
stems
probably
(eg Post
from
dependence
dependence
entrain
influence
peak of
previous
the
concentrate
food
of
diet
in
items
a major
the
the
observation
on vines
to
of
with
in
a peak
baboons
Foods Eaten
apparent
to
food
type
one
on the
effect
likely
had
Plant
climate
certain
turn
of
overabundance
the
the
in
which
is
of
abundance
a time,
seasonal
rainfall
in
on the
rainfall
strongly
peak
Differences
previous
comm)
pers
while
apparently
their
fruiting
and
in
the
an
ranging
season
7.13.4:
Relationship
The less
This
between Fruit
fruit
proportion
that
the
rest
in
was eaten
that
the
of foods other
than
fruit
diet
their
of
the
any month,
may indicate
relationship
small
and Seeds
may then
seeds
more
baboons
depend
seeds in
or
were
having
upon
their
be made up of
eaten.
a
diet,
and
fruit
or
either
seeds.
7.13.5:
Dependence on Trees
Dependence
to
lack
a
artificial
park
is
7.13.6:
grass
loss
of
item
with
data
is
individual
when the
food
terrestrial
boulders
was
were
accounted
such
of
item
by
under
in
direct
be
to
This
result,
of
Mandrillus
nemestrina,
survival
tend
studies
the
differential
or
categories.
in
if
smoothing
differential
remains,
useful
as Macaca
such
This
"fruit".
observation,
coarser
may
as
feeding
be pooled
can be pooled
they
because
occurs
as
thesis,
out
cancel
while
"difficult"
and
sphinx,
Discussion
Boulder-Rolling:
boulders
as part
leucophaeus.
There
major
considered
types
primates
Mandrillus
7.14:
types
is
this
season
cannot
remains,
classes
presumably
food
in
dry
is
the whole
since
season
the
of
observation
feeding
major
of
diet
used
by direct
classes
dry
there
Assirik
Mt.
and Feeding Remains
collected
major
At
the
may be related
seasons
1977).
and Verschuren
from
in
in
beginning
the
dry
plants.
cover
at
collected
considered
observability
not
year
(Dupuy
data
data
of
effect
ground-level
Observation
Direct
and in
sites
arid
forb-level
and
every
policy
While
of
of
burned
management
in
on trees
for
in
source
the
a marked
rolled
by
the
wet
of
seasonal
by
baboons
presence
Mt.
at
of
animals,
Animals
season.
animals
change
in
the
found
diet
7.31
in
the
frequency
This
Assirik.
mostly
beneath
of
insects,
boulders
baboons
at
with
was
which
probably
beneath
the
may be the
Mt.
Assirik,
in
especially
observed
the
profitable;
baboons,
Very
A variety
Baboons
did
animals,
basis
boulders,
data
of
the
boulders
to
roll
them and not
by
Fady
(1972)
too
beneath
them
beneath
the
could
how
boulders
in
how far
by
expectation
which
returned
beneath
the
of
of
selected
and
was
not
boulders
by
limit
The
highest
probable
work
baboons
that
boulders
were
(peanuts)
placed
tunnels
small
made
work
experimental
by strength
was determined
reward.
size
required
the
since
Further
food.
on the
Experimental
over
Some baboons
On
roll.
strength
rewards
of
few
were
to
limit
upper
case,
turn
to
the
there
which
returns.
the
not
efforts
out
swollen
injuries
by the
was set
is
upper
the
the
expectation
scoop
as
gripping
about
much effort
that
experimenter.
to
such
seen with
but
required
this
animals
these
of
for
find.
to
cautious
random.
expectation
the
protein
baboons
apparently
benefit
for
the
and
sampled
least
cost.
Damage to Plants
7.15:
The
three
A.
probably
dicitata
plants
digitata,
probably
both
swallowed
the
survived
the
and deposited
in
growth
out
of
to
new trees.
be
and
benefit
a net
of
spat
likely
most
B aethiopum,
gained
many flowers
since
to
the
boulders
clarify
much
were
by
at
rolled
great
in
them,
cause
possible
that
made very
for
massive
by the
of
sometimes
boulders,
baboons
suggests
he observed
that
is
sources
is
can be imagined.
which
it
the
that
the
causes
small
foraging
dangerous
were
boulders
ignore
these
of
but
of
easy
noticeably
frequently
so
form
particularly
not
Baboons
roll
large
and
were
possible
not
to
tending
size,
of
is
is
this
important
potentially
forearms,
painful
known.
the
baboons
that
are
such prey
boulders.
the
rolling
animals
hid
boulder-rolling
suggests
year
boulders
and the
obviously
of
perhaps
few
That
season.
or perhaps
scorpions,
or
wet
some times
at
highly
the
fruiting,
to
cheek
It
often
is
7.32
the
far
in
possible
baboons
from
places
that
seeds
the
of
parent
fruit
ripe
tree
apparently
the
A.
baboons,
the
of
actions
the
were
However,
erinaceous.
and
pouches
faeces,
P.
from
by
affected
baboons
and
suitable
of
Mt.
Assirik,
like
those
have been responsible
of A. digitata,
the
Each
which
P.
in
B.
field
beneath
plant
may depend
part
in the
and the
spread
L.
of
tree,
Such
stems
growth
Much of
be
cause
research
the
erinaceous
destruction
seedlings
newly
and
attentions
the
of
this
of
success
the
also
by burning
undergrowth,
trees
of
and
germinated,
baboons,
but
Oxytenanthera
help
the
on
population
bamboo,
with
was covered
limit
to
but
insubstantial
between
indicate
P. papio
baboons
are
that
its
and
on
culled
the
damage.
compare
damage done
the
the
park
each
but
also
consume trees.
several
mature
by
as
to maturity.
before
out
ecological
might
area
(such
trees.
is
this
of
parent
probably
impact
relationship
carried
their
up to 2
spent
survival
damage,
their
baboons.
dispersal
of
baboons
field
are
The
threatened
to
survive
here
and baboons
not
but
the
managed to
into
damage caused
by
eaten
damage from
from
and shoots,
bamboo,
of
presented
they
by
agents
distances
seen
should
destroy
long
frequently
research
that
the
seeds.
other
were
and many bamboo shoots
grounds
upon
seeds
By eating
environment
germination
avoiding
the
is
results
successful
reproductive
eating
principally
seemed insignificant.
further
serious
reproduction
vegetative
if
and
evidenced
whose
baboons
The
as
many of
windborne,
few and conspicuous,
are
depositing
abyssinica.
are
their
upon
flowers,
thousand
several
seeds
survived,
a single
One plant
such
and N. latifolia
area.
seeds
hours
P.
groves
plateaux
Their
may suffer
germinating
which
in
aethiopum
elephants)
The
many have
the
bears
destroyed.
are
but
species
tree
seed.
depends
baboons,
only
laterite
1979),
et al.
of the characteristic
M. alnifolia,
the
edges of
to
they
maturation
the
the establishment
erinaceous
survive
baboons
the
Ghana (Lieberman
in the woodland.
acida
Its
for
Shai Hills,
D. mespiliformis,
beneath
soil
the
of
B.
individuals,
may
easily
7.33
The annual
year.
aethiopum
palms
together
have
a
by baboons
I have
destroyed
with
far
any
greater
with
fires
not
seen many
by
fire;
damage
long-term
of
influence
on the
increase
the
expense
of
of
species
Data
in Diet
from
was based
this
plants.
The
by examining
may become
or
too
study
are
1978).
different
We
year.
of
may
identification
direct
and,
in
by the
the
former
this
study
on
direct
by
observing
While
seats,
of
examination
legitimate
some items
may be
faecal
samples
draw
to
feeding
others
general
some
compete
basis
this
chimpanzees
at
Mt.
Assirik.
foods,
as
the
seeds
such
apparently
unable
to
a higher
to
open.
biomass
of
Baboons
They
of
that
the
it
were
This
breadth
the
7.34.
capable
which
of
of
diet
chimpanzees.
and
the
same
at
diet
the
of
than
eating
chimpanzees
may have
of
baboons
chimpanzees
had a wider
also
times
seems unlikely
numbers
probably
aethiopum,
did
same foods
analysis
B.
than
rather
the
for
limiting
are
competition.
at
same foods,
different
tentatively,
rather
Mt. Assirik
eat many of the
probably,
general
On the
at
feeding
sustain
since
1953).
in
at Mt. Assirik
conclude,
not
year.
baboons
the
extents,
did
chimpanzees
times
1979)
sampling
for
Baboons and chimpanzees
to
and
(Lockte
seems
(Baldwin
however.
conclusions,
but
analysis
clear
It
park.
comparable,
not
only
diet
some food
of
Mt. Assirik
at
between
discovered
the
and Chimpanzees
strictly
not
the
at
alter
distribution
national
chimpanzees
relationship
or
this
sample
macerated
distinguishable
(Moreno-Black
on
scats
the
may also
species
Man may therefore
Baboons
study
Burning
cause.
fire-resistant
and
in
others
is
could
restricting
on faecal
largely
observation.
and
of
betwen
the
from
and data
scrub
encouraging
Overlap
baboons
the
of
by destroying
while
species
7.16:
proportion
other
baboons
than
park
helped
that
by
did
some
were
them
CHAPTER8: The Influence
of Food on Ranging and Troop Size
INTRODUCTION
8.1: Influence
The feeding
the ranging
their
Hamilton
strongly
P. ursinus
respectively.
to
habitat
the
test
ranging
large
the
the distribution
both
Rowell
or
grove
of
troops
of
5),
but
is
nothing
their
species
fig
trees,
is
known
In
food
plant
of
and
while
P. anubis
of
ranging.
of
(1966a)
by P. papio
ranging
into
and
responsive
about
this
more
I
chapter
influenced
the
at Mt. Assirik.
Troop
around
Size,
Mt.
Proportion
and the
Assirik,
(Chapter 3), at least
troops
tree
ranging
no single
of baboons
region
the
(Chapter
that
hypothesis
8.1.2: Home Range Area,
In
a fig
may influence
which
pattern
that
since
Thus
At Mt. Assirik,
types
factors
detailed
found
an insight
to provide
Assirik,
patterns.
influenced
fruiting,
gross
ranging
(1978)
may help
at Mt.
troops
of
may alter
et al.
of P. papio
behaviour
behaviour
food
on Ranging
of Food Species
in
of Fruit
P. papio
lived
in
Diet
the
exceptionally
large
one of which had an unusually
home
(Chapter
5).
range
Animals which depend largely
than do animals
that
suggests
will
have large
used to test
in larger
troops
baboon troops
which
home ranges.
this
that
prediction
(Milton
which are more foliverous
1971), and tend to live
This
tend to have larger
on fruit
Data from Mt. Assirik
baboons there
and Harvey 1977).
much fruit
diet
in their
on baboons may be
Data from the literature
hypothesis.
the
1976,
Schoener
May
and
(Clutton-Brock
include
home ranges
were used to test
depended particularly
strongly
the
upon
fruit.
8.1.3: Home Range Area, Troop
Clutton-Brock
food sources
home ranges.
clumped,
and
are best
This
Size,
(1978)
Harvey
or more dispersed,
argue
by large
exploited
hypothesis
and Clumping
would
than
that
ranges.
8.1
of Food
that
and dispersed
clumped
groups of animals
predict
of
and Abundance
that
other
P. papio's
baboons
with
having
food
large
was more
smaller
home
Alternatively,
abundant food.
the
large
This proposition
Iwamoto (1978)
Firstly,
been limited
feeding
by their
station
size
during
who showed that
of many troops
constant
Severe seasonality
imposed
alter
social
in the
the
(Hall
structure
of
food
5) is
to seasonal
the
increase
in
(1977),
by Dittus
became scarce,
the sizes
living
on a population
Inadequate
may influence
et
food
on a
group
group
while
seen
differences
a change
(Caraco
size
We may therefore
size
test
and
whether
(Chapter
Mt. Assirik
at
to
similar
can dramatically
1976),
al.
foraging
1978).
in the abundance
changes
at a
years
Only one troop,
1963a, Hamilton
to seasonal
either
a response
1972).
foraging
had
in numbers.
and Harvey
variable
seasonally
several
a study
when food
diminished.
(Jolly
for
a concomitant
may impose restraints
1975, Clutton-Brock
Wolf
with
drought,
increased
by aridity
distribution
food
of
comes from
evidence
of Macaca ssrui.oº
Macaca fuscata
of
to
two sets of data.
When the monkeys were supplied
markedly,
a severe
supply of refuse,
those
a population
quantities
rose
Further
be a response
might
is suggested by at least
showed that
large
with
density.
population
of P. papio
food supply.
and troop
population
troops
in distribution
of food or
of food.
METHODS
8.2: Influence
of a Single
Adansonia digitata
on baboon ranging
the
meant
and
diet
apparent
that
(3)
seen from
the
its
because (1)
location
of
up to tkm away.
distribution
trees
of
its
baboons
the
of
on Ranging
was chosen to examine the influence
conspicuousness
known A. digitata
the
Food Species
fruit
this
at
individuals
meant
portion
(14%) of
(2)
comparative
rarity
site;
the
on this
troop on two days when A. digitata
of
in
showing
could
these
was fruiting.
with
the
be plotted;
trees
could
distribution
the
the home range
map compared
8.2
its
species
baboons
A map was prepared
in one portion
trees
formed a major
of
that
of one species
of
of
one troop,
ranging
be
of
and
the
8.2.2: Influence
Each
of Distribution
the
of
characteristic
by
the
across
between
Table
7.4
habitat
types,
and
habitats
entered
using
to grow in those
habitats.
better
the
of
food
The measure
availability
within
for
the
Potential
diet
feed
sophisticated
of
of species
to
use
Influence
Troop
Size
Patch
types
were
photographs,
habitat
calculated,
to
size
of
diet"
it
would
also
and across
and the
measured
dispersion
by superimposing
the
and estimating
the
of
nearest
area
hectare.
and a mean patch
size
8.3
the
which
habitats,
free
of
each
habitat.
such
with
record
(A more
the
potentially
relative
of
patches
on
the
by contiguous
was estimated
modal
Ranging
patch
by dividing
and
habitat
various
on a map drawn
and
of the
these habitats.
Habitats
of
in
available
the rank order
contaning
a grid
Median
ranging
)
tau, with
covered
food
of
of each species
account
of the diet
and Dispersion
of the
same habitat.
the
A
with
to the feeding
habitats.
quadrats
a rough
habitat.
in different
into
take
only
frequencies
from
known
species
examine
available
contributed
species
(Chapter
estimate
measure
with
the diet
in
A
in
overlap
that
by summing the contribution
other
Clumping
the
species
was compared, using Kendall's
8.2.3:
in
we shall
circular.
with which baboons entered
frequency
is
light
The rank order of the proportion
each habitat
any habitat
importance
different
which
within
food
whose
be
various
available
of
habitats
as an independent
invalid
then
measure
abundances
in
was established
contribution
in
species
frequency
the
of
a
food
degree
number of
by their
be used
on the
the
the
food
of
"potential
the
order
is
were described
of
the
2
Chapter
habitats
table
the
found
species
to
by the proportion
weighted
the
is
would
reasoning
is
tau, with
therefore
seen to
circularity
the
in
distribution
containing
habitats,
is
It
were
this
amount
weights
baboons.
baboons
from
species
potential
measure
behaviour.
the
The rank
Kendall's
The number of
described
shows
quadrats
on Ranging
Some of these
was calculated.
5) was compared,
measure
types
of plants.
(1979).
baboons
which
habitat
seven
community
Baldwin
of Food Species
from
aerial
areas
size
the
the
of
were
area
of
Figure 8.0:
between clumps
of mean distance
calculation
of
a -habitat,
given mean clump size and number of clumps in known area.
d= distance
between nearest
points
of adjacent
clumps
between centres
s= distance
of adjacent
clumps
a= mean area of clump
(calculated
from area)
mean
r=
radius
of clump
A= inverse
of clumps
of density
°
°°°°
00000
00
°0000
°area
a°
-'
v v0
00ä
area A
r
d
sh
.13
b/2
each habitat
in the field
The mean distance
habitat
area by the number of patches
between the
nearest
were not measured directly,
Assume regular,
close
Then the distance
d
of
adjacent
but were estimated
(Fig
of the patches
packing
habitat.
patches
of
a
as follows:
8.0).
= d, where
we require
2(h
points
of that
-r)
but
r = square
(a/pi
root
)
and
D = 1/A
u/(
2hb)
1/(
(sqrt
h=
sqrt
(1/(
d.
2x
3) xh squared )
therefore
(sgrt3)
)
xD
and
Habitat
( (sqrt(
for
maps prepared
apparently
(sgrt3)
1/(
Dunbar 1974c, Harding
1976, Moreno-Black
1966a, ) and cannot be used for
carried
out
Dunbar
kindly
Assirik.
research,
in
the
home range
size
between
and
troop
site
could
as for
the
clumping
then
et al. 1971
photo
and inter-clump
size
large-scale
size
The map of the
study.
from an aerial
Mean clump
this
this
where Aldrich-Blake
was prepared
same way for
The relationship
with
in Ethiopia,
gave me access.
calculated
were
the
their
1971, Dunbar and
al.
et
on baboons
studies
other
Nagel
1973,
Rowell
1977,
Maples
and
comparison
in the Awash valley
study site
in
publication
(eg Aldrich-Blake
much detail
omit
)-r)))
xD
area
to which
distances
of the habitat
be examined
Mt.
around
across
and
two
troops.
8.2.4: Home Range Area and Proportion
Data from the literature
between the proportion
troops
of various
and proportion
way that
species
of fruit
obscured
of
of the Diet Consisting
was also
fruit
in
the
used to examine the
diet
might both be affected
the relationship
8.4
correlation
and the home range area of
of baboons of the genus Papio.
available
of Fruit
between them.
Home range area
by rainfall
A partial
in a
correlation
Hap 8.1:
full-day
two
Part
of
of known Adansonia
ranges
digitata
plotted
trees.
relative
to
distribution
the
"N
T
KEY:
digitata
Adansonia
two
day
ranges
tree
ý'--ý
...........
cliff
line
t"
ýI
01
I
ýý1
ti
"r
?1
Ja
r7
.'i
,"'ý
.s
1
11
"1
"
"Y
mot.
.40
00
e
.0p
I kilometre
between home range area and proportion
for annual rainfall,
was also carried
of fruit
in
the diet,
correcting
out.
RESULTS
8.3: Influence
of a Single
The locations
of
the
of
portion
ranges
indicated
the
that
tracks
exact
path
end in
into
in
the
the
question
to test
giving
kilometre
of transect.
which 21 trees
troop.
always
trees
Map 8.1.
fissions
Poor
that
of this
the
of
trees
per
troop
meant
these
they
the
cases
seen to
were
climb
They
visited.
days.
24km of 200 metre-
result,
the area and 29 trees
0.17
Parts
visibility
known; in
trees
south-eastern
in the area on one or both
the significance
of
the
in
The baboons
map.
of 45 A. digitata
a density
survey.
in this
counted
sq km, or
1.2 trees
per
Roughly 9.5 km of day range is shown on Map 8.1, in
by the main group on each day range.
were visited
significantly
visited
was not
were drawn through
wide transects
the
in
plotted
Temporary
of
on the
29 out of 45 A. digitata
In order
are
plotted.
troop
trees
troop
path
marks
from 25 out
and feed
visited
df
of
Fable's
of
are also
by forks
on Ranging
known A. digitata
all
home range
two full-day
are
of
Species
trees
more
than
expected
(chi
They
squared = 7.897;
= 1; p<0.01).
8.3.2: Food Availability
The rank order
enter
rank
quadrats
order
(Kendall's
diet
of
and Use of Home Range
of the
frequencies
containing
the
numbers of
food
various
with
which the baboons tended to
habitats
species
available
within
contained
tau =0.24; ns. ), nor as the rank order
potentially
was not
from each habitat
2).
in
(Plate
1.1).
The largest
contiguous
8.5
those
habitats
of the
tau = 0.14; ns. ).
(Kendall's
in the aerial
were distributed
same as the
of the proportion
8.3.3: Clumping and Dispersal of Habitats
Seven habitats
were distinguished
These habitats
the
a mosaic
photograph
over
area of any habitat
the
(Chapter
field
in the field
area
area
was one of 379ha of closed
woodland , while the modal contiguous area for
Median patch sizes are
was less than one hectare (Table 8.1).
each habitat
also
shown in this
Table 8.1:
Central
Table.
Tendency in Patch Size
Habitat
6.00
3.18
2.15
1.41
Grass
on laterite
Combretum scrub
Scrub
8.3.4:
Grassland
patch
single
1158ha in
of
field
the
field
0.97
<1
5.2
275
1.9
0.88
<1
13.4
166
8.1
0.32
<1
3.2
502
0.6
of
and p=0.46
We should
are roughly
kilometres
patches
(this
was high,
in
(Table
that
predict
group
troop
than
it
to compare mean monthly
8.6
studies
boundary
between
(median
the
test
p
and home range
areas
areas
were 44
and the troop
sizes
250 and
et al.
1971).
Size
was higher
was in
the
The home range
square,
on Troop
both
sizes
and Aldrich-Blake
size
beyond
over
8.2).
at the two sites.
study
The largest
distances
mean
and
sizes
area
extending
scrub-grassland
were comparable
of Food Abundance
Mean foraging
valuable
clump
of
(1971).
et al.
an unknown area
over
square and 4.3 kilometres
Effect
abundance
area
from the
those
were compared with
respectively).
comparable
87 respectively
8.3.5:
Median
therefore
and Ranging and Group Size
of Habitat,
(and
area
area).
points
nearest
= 0.5
<1
<1
<1
by Aldrich-Blake
studied
A/n
(ha)
5.7
12.2
7.1
12.4
was a contiguous
their
Area
No. of
(sq km) patches
An
7
218
262
181
from Mt. Assirik
used by the troops
at Mt. Assirik
0.4
26.6
18.7
22.4
Clumping and Dispersion
The data
Mode
Median
Gallery Forest
Open Woods
Closed Woods
Scrub
in Habitats
the
foraging
in
the
dry
wet season,
season.
group size
with
It
when food
would
rainfall
be
for
Table 8.2:
Clumping and Dispersion
Habitat
of Habitats
n of
patches
at Two Sites
mean of
patches
per sq km
mean area
of patch
(hectares)
distance
between
patches
(m)
Mt. Assirik:
7
218
0.08
2.42
5.7
12.2
5103
583
262
2.91
7.1
590
Scrub
181
2.01
12.4
675
Grass on Laterite
Combretum Scrub
275
166
3.06
1.84
1.9
713
799
Scrub Grassland
502
5.58
8.1
0.6
Riverine Forest
Open Woods
1
0
0.06
0.0
Closed
0
0.0
Gallery Forest
Open Woods
Woods
Closed
556
Awash:
Woods
Scrub
that
0.0
Combretum Scrub
0
0.0
Scrub Grassland
Grass
82
43
4.62
2.42
apparent
party
accurately
P. ursinus
5 troops
of
were used to
test
from
and the proportion
the
tau
of
their
diets
was a non-significant
home range
= -0.25;
and
p=0.38),
the
the
which
--
in
could
in this
of
not be estimated
study.
Made up by Fruit
and 1 of
P. cynocephalus,
between home range
fruiting
fruit
trees
between
relationship
became stronger
8.7
size
2 of
found
283
787
14.1
2.8
relationship
negative
proportion
--
of Diet
P. anubis,
360
2.9
sample sizes
Home Range Area and Proportion
Data
There
the monthly
enough with
--
7.55
0
but mean monthly
-
--
Grass on laterite
month,
8.3.6:
134
101.0
in
the
when the
diet
(Table
the
area
8.3).
size
of
(Kendall's
correlation
was
corrected
for
(partial
annual rainfall
= 0.53;
correlation
Table 8.3:
Home Range Area and Proportion
of 8 troops
of baboons of
% Fruit
Home Range
Species
in the
of Fruit
the genus Papio
P. anubis
0.90
40.8
P. anubis
1.12
82.7
P. anubis
P. ursinus
1.12
6.7
48.6
77.0
Source
Dunbar and Dunbar 1974o
Dunbar
1974o
and Dunbar
Ransom 1971; Oliver unpub.
Hamilton et al. 1978 (K)
19.7
9.5
Harding 1976
P. cynocephalus
P. cynocephalus
40.2
61.0*
27.1
15.5
Post 1978
Oliver
unpubl.
P. anubis
52.0
46.0
Popp 1978
P. anubis
Diet
Note 1: (K) = Kuiseb
'Oliver
2:
used 1 km. sq. quadrats
(Ruaha)
this
to calculate
tend to give an
Dunbar (in prep) has shown that large
figure.
quadrats
He provides
when
which,
a correction,
of the true area.
over-estimate
42
Oliver's
to
estimate,
gives
sq. km.
applied
60.3%
up
of
makes
(Kendall's
Since
diet;
home range
tau = -0.14;
p_0.6;
the relationship
and home range area did
cannot
be attributed
area
partial
km)
44
were
sq
=
the large
not hold,
the
high
proportion
(fruit
included
p=0.23).
= -0.31;
correlation
between the proportion
to
P. papio
became worse when data for
In both cases the correlation
in the diet
of fruit
home range area of P. papio
of
in
fruit
its
apparent
diet.
DISCUSSION
8.4:
Influence
of Distribution
The location
of
baboons at
therefore
distribution
be
of food
of Food on Ranging
trees
Mt. Assirik.
couched
of
food.
at
influences
the day ranging
Any explanation
least
With
partly
appropriate
8.8
in
of
ranging
terms
of
data,
the
of one troop
must
patterns
response
rank
to
the
order
of
preference
with
habitats
be compared
could
diet
potentially
each
quadrat
data
presented
nor
Food
influencing
behaviour,
social
fragments
proportion
troop
of
the
food
of
the
diet
the
by areas
sites,
in
those
factor
only
suitable
for
in
which
areas
or meet other
not
different
found
not
feeding
to detect
likely
does
in
potentially
into
entry
However,
found
probably
between
of
home range
species
the
of
analysis.
be affected
may also
are
this
various
proportion
Frequency
for
therefore
routes
the
with
use
of
is
efficient
the
of
the
the
containing
habitat.
study
number
which
ranging,
month
each
that
availability
quadrats
by
this
the
the
with
in
shows
with
closely
habitats.
from
low
here
entered
month
available
was too
correspond
habitats,
baboons
which
fragments,
and
depends
upon
so on.
8.4.2:
Clumping
The size
level
the
which
areas
of
probably
the
which
analysis
of
type,
habitat
consider
in
in
differences
research
by
survey,
and
areas.
If
noting
two troops
same field
sampled
area
at
yet
help
of
habitat
to
explain
types
This
themselves
Mt.
at
Assirik
intensively
accurately
several
coarser
large
the
for
Clumping
home ranges
attempt
and dispersal
where
the
responsible
in
these
by
of
link
clumping
in
range
food
problems
those
be found
could
to
the
ecological
baboons
each
widely
Future
detailed
areas
in
be
sites.
with
the
of
quadrats
the
a
habitat
of
confounded
two
cope
would
habitat,
of
baboons
is
home range
in
to
of
these
behaviour
level
8.9
troops
in
analysis
dispersal
analysis
areas
dispersed
does not.
two
the
probably
areas
appear
might
used
different
with
dispersal.
in
than
and
do not
photos
finer
rather
clumping
areas
habitats
a slightly
food
with
site,
the
Africa.
topic
small,
selecting
the
in
the
on this
aerial
home range
locations
separated
in
species,
are
patches
though
The largest
food.
the
of
here,
with
food
of
from
distribution
a "patch"
be considered
Differences
differences
for
the
as dealt
stands
identifiable
types
can be considered
realistically
could
be patches.
to
of Habitats
an area
of
of
areas
and Dispersal
at
be
could
home range
species
may
and dispersal
8.4.3: Fruit
and Home Range Area
There was a non-significant
the proportion
Assirik
good explanation
It
diet
that
home
of
ecological
analysis
behavioural
to test
on which
animals
Baboons,
because
further
but had large
home
not a
fruit
of
test
such
are
requires
prediction
in
unattempted
home range
probably
that
areas
baboon
large
are
best
the
not
in
be a strong
would
a
as yet
their
primates,
of fruit
of the proportion
detail
of
with
most other
with
Data from Mt.
of baboons is therefore
sources
To
area.
a fineness
at
ecology.
by comparison
the
of
range
diet
a combination
dispersal
predictor
apparent
to the
in home range area of baboon troops.
of variability
the
and
still
when
contrary
(1971).
trend
in the diet
of fruit
seems plausible
the
of
is
which
and Schoener
in their
much fruit
included
home ranges to be smaller
was higher,
significance
The proportion
ranges.
for
(1976)
May
and
the
decreased
these troops
the
in the diet
of fruit
of Milton
predictions
trend
group
of
such a relationship.
8.4.4: Ranging and Mental Maps
If
day
forming
currently
species
corresponds
range
closely
the bulk
baboons must possess a mental
1970).
Altmann
featureless
scrub,
over
on
two kilometres
waterhole
an
a heavily
from their
as seems probable,
the
the existence
for
study,
hour
food
home range area (Altmann
for
evidence
in the course of this
moved for
troop
the
of
map of their
Circumstantial
arose repeatedly
diet,
the
of
distribution
the
with
in
one
and
of such a map
example; on one occasion
direction
overcast
day,
starting
place.
flat,
through
before
a
at
arriving
8.4.5: Use of Home Range
food
If
discriminate
are feeding
In
closed
alone
were
two habitats
on occurs
at a higher
found
woodland
tree
is
alone
between
most
particular,
were probably
in
availability
food
the
whenever
in
species
found
in greater
density
in closed
tended
be closer
considered,
to
the
baboons
8.10
than
since
woodland
have
If
should
which
woodlands
the
tree-based
been
they
in the other.
open and closed
together.
might
food
principal
in one habitat
density
baboons
the
considered,
expected
trees
foods
to
discriminate
by the
of
in
favour
analysis
the
food
at Mt.
(Byrne
These foods
may have found
in
its
altering
behaviour
part
P. papio
which
diet,
from Mt. Assirik
Foraging
of Optimal
2,
in
to
may have
Visibility).
and the
record,
open woodland
than
benefit
of
to test
Theory
to
responds
degree
the
and
to
are
alter
environment
alters
its
foraging
brought
out
by using
it
both
its
in
changes
which
two specific
foraging
of optimal
predictions
These are:
(a) Abundance of Food and Quality
Cody (1974),
Firstly,
Pyke et
(1977)
al.
up
the
so
much less
than
respond
the baboons
Ellis
and Schoener
quality
to changes
only
applies
the
in food
at Mt. Assirik
Secondly,
abundance once a threshold
Data
from
restricts
low abundance the
Mt. Assirik
place
this
prediction
over
a timescale
Animal
animal.
populations
in numbers. ) Data from
prediction.
goes down.
that
as food abundance goes
animal
Westoby (1974)
By contrast,
of a food in the diet
changing
of
the proportion
take
by an increase
also predict
that
times
the
up.
(This
of Diet
of the diet
only
of
that
goes
that
can be used to test
predicts
availability
eaten
(1976),
Dunham
and
as food abundance
Estabrook
predict
abundance
changes
the above authors
up, so the diversity
all
food
time
generation
(1976),
et al.
(1971)
the
of
to longer-term
of Diet
(b) Abundance of Food and Diversity
that
foods
Chapter
diet
research
habituated
well
forb-level
their
of
future
for
in the feeding
represented
a larger
basis
(see
woodland
open
under
as expectations
theory.
might
However,
or absence
on presence
now reasonably
comm.
the
Predictions
The manner
goes
are
the
indicated
here.
8.4.6: Testing
data
in
were probably
is apparent
by
baboons
pers
form
well
only
is
than
more strongly
depends
which
could
the
since
densely
more
baboons
This
species.
human followers
grown
here,
presented
Assirik,
woodland
of closed
will
not
of abundance is reached.
consumption
will
can be used
to
predictions.
8.11
include
test
when it
is
change with
He suggests
low.
fewer
items
between
these
Thus at
in its
diet.
conflicting
There is
roots
they
which
different
in
parts
quality
the
and P. cynocephalus
troops,
in the dry season but
(Altmann
available
rich
foods
foods
two sites
flushing
to
grass
grain
fed
are
such as leaves,
to
in
the
include
only
both
Finally,
Macaca
wild
and corms
becomes
when energyfuscata
the diet
from
high-
P. anubis
when it
or to fruit
dropped
are
eat
depend upon roots
1976).
1970, Harding
and Altmann
T. gelada
changes
comes from
evidence
in
to
to
seeking
in
and energy
that
showed
response
parts,
Further
switch
in energy,
in
plant
which
much time
spend
(1977a)
various
as cereal
such
poorer
grass
diet.
their
to
Dunbar
the
of
quality
nutritional
1976).
of
parts
of the food
else, ignoring
they eat almost nothing
have
would
(Wrangham
excavating
so the quality
in the dry season, but when in the
and roots
eat rhizomes
wet season green grass is available,
the
to support
primates
also increases.
in the diet
T. gelada
on other
as food abundance increases
that
the prediction
included
from data collected
evidence
troops,
(Iwamoto
1978).
There is
also
increases
abundance
dietary
so
(Pan
troglodytes)
chimpanzees
food
where
dwelling
(Suzuki
to support
some evidence
ate
Pan troglodytes
in
decreases.
diversity
as food
For
example,
in woodland savanna in west Tanzania,
living
was scarce,
that
the prediction
a wider
of
variety
Uganda, where food
did
than
food
was relatively
forestabundant
1969,1979).
METHODS
8.5: Quality
of Food
The data from direct
were
leaves,
flushing
twigs,
rich
into
arranged
roots,
in
16 broad
mature
fats
compounds and relatively
foods.
leaves,
Gum is
difficult
categories
shoots,
or
sugars,
easy of
to
but
bark,
8.12
flowers,
fruit,
seeds,
stems, base of stems, bulbs,
Those foods which were probably
low in
probably
access
categorise,
by baboons at Mt. Assirik
(buds,
and "other").
gum, animals,
proteins,
on feeding
observation
were regarded
since
it
is
secondary
as high
sometimes
toxic
quality
rich
in
but
carbohydrates
toxins
other
depending
and Bearce 1976).
feeding
of
food.
on the species
Since it
concentrations
from which it
was only rarely
classed
low-quality
as
foods.
flowers,
flower
quality"
foods, by contrast
buds, flushing
Thus
leaves,
as a low-quality
was soft,
were classed
fruit,
animals,
and bulbs
the other
with
it
classed
which seemed to be difficult
roots,
and
(Hausfater
is derived
Bulbs, eaten in the wet season when the soil
foods, while
tannins
of
(12 out of 2607 records
eaten
I have tentatively
or foraging),
as high-quality
I
high
sometimes contains
to dig up,
seeds,
were defined
shoots,
as "high-
which were defined
categories
as "low-quality".
8.5.2: Measures
of Abundance
No direct
measures
measures
therefore
were
abundance
Mt. Assirik.
With
that
Secondly,
baboons,
foraging
search
to,
gave rise
proportion
food
dry
abundance;
is
fruiting,
and
availability
season
Hall
social
in
fed.
here
food
of
flowering
the
These
data
to
provide
of foods
for
were
an
the
collected
indication
the baboons.
day.
here
a study
by S. Hall
abundance
and
1981).
The
as an index
less
of
of
social
in
food
the
in the wet season.
of
the
leafing,
from
and others,
seasonal
spent
on primates
plants
of
showed
and probably
showed
than
that
time
the
studies
abundance
species
of
of
1970, Post
used
in the dry
twenty
of
8.13
4
wet,
times
(1978)
food
baboons
came from
abundance
cycles
Loy
is
activity
the
low
of
1963;
since
than
Other
times
food
with,
coincided
behaviour.
of
implies
in
at
times
and Lee
an increase
season
assumed to have been lower
measure
analysed
(eg
in
apparently
Oliver
between
in
spent
dry
A further
baboons
social
specifically,
in the
season
in
dry
abundance
profitable
this
food
Park in Kenya.
plant
that
less
baboons,
the
activity
time
into
primates,
in
National
predicted
that
showed
of monthly
activities.
relationship
social
of
behaviour
food
a reduction
in
reduction
be extended
on social
of
a similar
show
(1971)
of P. cynocephalus
a troop
in
moving
as a measure
some other
and with
in Amboseli
rainfall
Schoener
would
encroaches
for
with
(1978)
-Post
indirect
Three
were available.
Firstly
used rainfall
foraging
shortage,
abundance
used.
was correlated
I have therefore
food
of
changes
which
and
are
in
the
8.5.3: Tests of Quality
of Diet
The prediction
concentrate
of
that
abundant
on high-quality
foods
high-quality
The proportion
foods
of Diversity
the number of different
food
the
number of species
in that
month's
The prediction
in
diet
season
in
season,
this
than
comparison
with
that
between
the
since
analysis,
that
August
in
would
August
the
have been with
feeding
previous
month's
size
between
of
baboons
each month's
sample
line
line
regression
not
could
began
in
feeding
the
two
size
in
a more diverse
ate
regression
study
monthly
in
and September
feeding
measures
included
influence
If
between
the
sample
the
of
for
the
the
and the
as the abundance
fell
of
were
in
record.
observations
relationship
seasons.
elevation
influence
was examined.
diversity
removing
of
the
The transition
season.
the
difference
wet and dry
be higher
would
months
feeding
month's
of the diet
items
of
effect
between
dry
the
record
the
of
the
on
had the
comparison
feeding
by examining
number
was regressed
months.
were
squared).
across
was plotted
feeding
of
had not been included
which
This
sample
measures
the
record
record
the
and two
Firstly,
feeding
month's
baboons
(chi
seasons
in that
size
the diversity
that
went up was tested
record.
that
in any month might
included
items
between
abundance
diet
of feeding
The relationship
of
the
proportion
to
of Diet
The number of observations
of
in
the
the
which
across
baboons
with
record
with
the
rainfall.
and compared with
food
feeding
was compared
food
allowed
by comparing
was tested
The frequency
low-quality
of
resources
monthly
tau.
seen to eat high-quality
8.5.4: Tests
the
Kendall's
using
rainfall,
in
foods
food
for
the
dry
for
the
wet
be included
September,
a month nearly
a
and
in
a year
the
future.
(1969)
Pielou's
Secondly,
modification
used to examine the heterogeneity
Pielou's
year's diet.
Diversity
where
month's
n(i)
is
feeding
of each month's.
(1912)
formula
each season's,
was
and the
index is:
SIGMA ((n1
=1-
the
of Ginni's
number
record,
of
and
x (ni -1)/(Nx(
observations
N is
the
8.14
of
total
the
N-1 ))
ith
number
food
of
type
in
observations
that
of
4
8.1:
Figure
Percent
of
(histogram)
4J
C)
.,.4
low
quality
food
plotted
(graph)
with
rainfall
20
0
0
W
cl
CD'
300
10
0
200
a
w
0
+1
0
C)
v
a
100
ý"
5
51
NDJFMAMJJ
450
Figure
Number
plotted
8.2:
feeding
in
items
food
different
monthly
of
for
that
month
size
against
sample
record
O
.ý
30
U)
'i
"e
.
4
GL
20
.O
O
4-4
o
10
"
aE
Wet
Season
Dry
Season
z0
100
Sample
Size
200
for
Month
9
feeding
for
increase
that
in
Thus if
10 observations
the
There
index,
of
like
food
index
and
partial
feeding
in
was therefore
size,
0.97,
index
if
10
the
index
the
this
index,
is
becomes
and Pielou's
to the number
sensitive
between
by
confounded
against
is
while
Any correlation
be
one of which
9 times,
for
Pielou's
A
size.
sample
for
controlling
rainfall,
(a)
types
importance.
relative
of
one eaten
therefore
food
9 species,
index
1974).
1 as the
their
include
significance
Pielou's
of
in
of heterogeneity,
might
rainfall
correlation
sample
for
(Peet
diet
towards
an
2 species,
indices
the
has
record
only
most other
types
tends
become similar
no known test
is
index
are made which
included
observations
0.2.
and (b)
number
twice,
eaten
This
month.
computed.
RESULTS
8.6: Testing
Predictions
The data
Foraging
of Optimal
by direct
collected
Theory
observation
by baboons
feeding
of
are
given in Table 7.9.
The
proportion
high-quality
of
negatively
correlated
p<0.001),
and the proportion
with
in
foods
feeding
the
(Kendall's
across months
rainfall
0.82;
=
tau
in the wet
foods was higher
of low-quality
was
record
season than in the dry season (Table 8.4).
The
and that
record
The
four
sample
the
the
variety
when the
effect
month's
(Kendall's
tau
0.56;
=
above
is
statistically
is,
within
each
items
of
in
sample
the
size
wet
feeding
month
the
season
was controlled
8.15
feeding
than
it
for.
(Fig
data
from
record
did
was
8.2).
line
regression
(Fisher's
significant
feeding
record
p<0.01)
are
p<0.01)
the
8.1.
Fig
least-squares
the
= 0.79;
trend
of
are
given
(r(p)
data
shown in
any
lying
points
that
p=0.030);
size
in
months
in
foods
low-quality
of
across
items
with
of
proportion
rainfall
of
This
season.
wider
of
through
plotted
the
number
correlated
All
in
variation
in
the
Exact;
contained
the
wet
dry
a
season
Figure
8.. 3:
feeding
in each month's
feeding
record
month's
between
successive
'size
items
Number
of food
in
previous
not
were
in sample
difference
20-
V
9O
rl
U
ý
NO
"0
>s
.0
.
w.
ýO
W
10
record
plotted
months
y
$ea5oý`
D'
®
`ýoý
ý'
ý`'o
O
-150
-100
-50
0
in Sample Size
Difference
This
Month's
and Previous
Feeding
Record
50
Between
Month's
100
that
against
Table 8.4:
Table for Numbers of Observations
of Feeding
Foods in Wet and Dry Seasons
on Low-Quality
Contingency
(Low-quality
foods defined as leaves, bark,
twigs, roots,
gum)
stem, base of stem,
Season
Wet
Dry
127
102.3
84
108.7
Observed
Expected
chi
8.6.2:
Change in Diet
The signed
with
Time
difference
in
record
between
size")
was calculated
included
not
were
is
items")
each pair
food
The number of
= 11.576;
squared
of
and is
items
in
the
number of
in
entries
in
previous
each month's
between August and September is not plotted
for
sample
of Fig 8.3.
feeding
record
that
record
("new
food
(The datum for
axis.
feeding
in
axis
feeding
month's
the
("difference
months
on the horizontal
plotted
on the vertical
plotted
= 1; p<0.001
consecutive
included
the
df
the transition
in the
reasons explained
Methods. )
There
was a positive
new items
the
number of
size
between
the
(r(p)
(wet)
= 0.94;
regression
season,
not
t=
between
were
10.9;
p<0.001;
sample
(for
both
the
each
number
on the
season,
t=4.89;
in
(dry
zero
but
p<0.02),
in
two
the
of
the
sample
were stronger
from
Thus
other.
new items
of
slopes
different
season,
from
seasons
The
between
p<0.05)
difference
and the
significantly
were
relationship
each month's
feeding
season.
of the seasonal
dry
= 0.78;
within
= 0.98).
wet
and the
dependent
The elevations
different
(dry)
different
size
diet
any month's
The correlations
r(p)
lines
was not
record
in
months.
significantly
(r(p)
correlation
regression
intercept=
8.16
lines
8.83±0.98
were significantly
(95% confidence
limits);
for
the wet season,
irrespective
in
eaten
five
of sample size
each month in
intercept
= 14.77±3.89)
or six
(Fig
more food items
8.3).
Thus
were seen to be
the wet season than were seen in
each month in
the dry season.
8.6.3:
Heterogeneity
Pielou's
Assirik
is
(0.92)
index
0.96
seasons
rainfall
for
the
(Table
8.5).
are
p=0.26),
between
diversity
the
feeding
The index
Pielou's
is controlled
so that
in
Record
year's
similar.
when sample size
df = 9;
Table
of the Feeding
this
feeding
index
record
record
for
the
index
is
fails
correlated
correlation
to
and food
(0.95)
wet
not
(partial
for
baboons
of
abundance.
Pielou's
Index of Heterogeneity
for Plant Food Types by Month
Dry Season
i
Nov
Dec
Jan
Feb
Mar
Apr
Month
i
0.723
0.663
0.825
0.928
0.872
0.921
Season
i
Aug
Sep
Oct
0.924
Wet Season
Month
10.866
Season
I
Year
Jun
Jul
0.879
0.919
0.948
0.961
8.17
0.891
and dry
with
= 0.37;
show any relationship
8.5:
May
Mt.
at
0.876
0.859
8.7:
Figure
Monthly
variation
19
from
species
monitored
in number of
fruiting
whose
food items
different
available
cycles
were
and leafing
20
C)
w
0
A
10
z
NDJFMAMJJASO
Dry
Season
Wet Season
8.6.4:
Seasonal Availability
The baboons
ate
parts
and fruiting
flowering,
of Food
19 of
of
the
20
were monitored
cycles
whose leafing,
species
(Table
were eaten
8.6).
Table 8.6:
Food Items Monitored
Through Year
Flush
Species
Flowers
digitata
Adansonia
Afzelia
africana
Bombax costatum
Ceiba pentandra
Cola cordifolia
Diospyros
mespiliformis
Erythrophloem
suaveolens
Ficus leucardii
Ficus
Seeds
Fruit
Shoots
ft
ft
*
umbellata
Grewia lasiodiscus
Hexalobus
monopetalus
Khaya senegalensis
heudelotti
Landolphia
Nauclea latifolia
Oxytenanthera
abyssinica
Parkia
biglobosa
Pterocarpus
erinaceous
Saba senegalensis
Spondias mombin
Strychnos
spinosa
Vitex
madiensis
*
*
*
sp. of Sapotaceae family
The number of
wet
edible
by month
month
plotted
in
(medians test,
This
have been a wider
variety
Data
from
relationship
annual
rainfall
14
other
troops
items
were
dry
season
the
is
in
the
available
(median = 9.75)
sample indicates
available
19 species
that
there
may
to the baboons in the wet
species
of
baboons
between the proportion
(Table
these
in the dry season.
season than was available
Comparison with
small
foods
of
More
in
than
p=0.004).
from
available
8.7.
Fig
(median = 16.5)
season
8.6.5:
items
8.7).
can
be
of high-quality
Rainfall
8.18
examine
the
food in the diet
and
used
was estimated
to
in
by
cases
some
figures
using
published
Table
8.7:
Proportion
animals)
from
nearby
Food (fruit,
1 Populations
of High-Quality
in the Diet of
% high
Species
Rainfall
Source
(mm)
Quality
14.4
P. anubis
flowers
seeds
of Baboons
Harding 1976
Popp 1978
Dunbar and Dunbar 1974c
595
46.9
56.4
1099
1105*
65.2
83.2
1380*
Ransom 1971
Dunbar and Dunbar 1974c
Oliver pers comm
P. cynocephalus
25.3
1105*
1679
P. ursinus
32.2
74.6
225
18*
Post 1978
Hamilton
et al.
P. hamadryas
66.0
Kummer 1968a
P. papio
T. gelada
84.3
21.4
665*
,941*
1385
This study
Dunbar and Dunbar
7.8
7.5
1476
1102
Iwamoto pers comm
Dunbar and Dunbar 1974c*
*
These figures
stations
meteorological
1975
from nearby meteorological
are estimates
(Dunbar
1976
and Sharman in
stations
No relationship
prep).
was found.
8.6.6:
Diversity
Quality,
At times
for
record,
abundance,
any given
sample
rate
The diversity
months.
Pielou's
at
food items
food,
but to
food
items.
Adansonia
in the first
of
within
some other
For
digitata)
half
index,
that
the
each month is
variable,
example,
feeding
upon which
8.19
feeding
was in the dry season
record,
measured
relative
emphasis on different
not
one
food
to total
abundance of
type
This
need not imply
by
abundance of
(the
the baboons depended heavily
of the dry season.
the
but was not correlated
related
was
in
was high,
such as relative
there
abundance was low.
included
were
each month's
may imply
This
plant
new items
of Ginni's
modification
rainfall.
when food
than
size,
which
in the feeding
more items were included
in the wet season months than it
was higher
record
with
of high
the
Similarly,
and Abundance
single
fruit
(Table
that
of
7.9)
food,
in
fruit,
the form of A. digitata
this
type
food
imply
that
other
food type at that
When food
to
the
(1974).
is
than
food
amount of
than
by time
to
true
for
baboons
searching
in
decrease
the
two further
dry
the
Firstly,
dry
and
of
gait
is
reason
in
which
assumes
that
diversity
environments,
invalid;
there
would
not
and
that
the
of
low
There
are
times
model.
the
feeding
such
as
of
in my sample
importance
season
than
the
feeding
record
examine
this
and
will
the
value
a wider
variety
8.20
is
foods
In
this
of
the
currently
of
not.
Mt. Assirik,
in
the
grasses,
diet;
the
of
there
for
season.
of
would
dry
this
be a
season.
possibility.
it
as
(eg
more items
wet,
wet
and there
diet,
of
If
in
dry
pause
and so on).
abundance
against
no discernible
or
theory
biassed
the
fungi;
to
the
in
diet
the
proportion
available
probably
results
account
at
were
a major
foraging
that
the
between
changed
for
in
both.
foods
may have
many of
be needed
optimal
is
rather
moving
unlikely
of
is
a large
season,
in
to
when
certainly
diversity
decreased
them were of
dry
diversity
while
of
(1974)
represented
ground;
the
in
low
Secondly,
Westoby's
little
with
that
on the
research
record
true
time
spent
were
feeding
diversity
the
were poorly
eaten
misleadingly
Further
in
believe
seeds
were
nature
such
many populations
insects;
dry
seems
a way that
be eaten
could
to
the
diet
the
of
nature
in
a baboon
quality
is
almost
it
of
explanations
an increase
that
by
for
in
Thus
food
that
diet
due
by digestion
baboons
of
prediction,
is
the
of
counter
season.
the
seasons
recording
Items
diversity
plausible
in
4).
run
so diversity
assumption
the
therefore
assumption
limited
is
where,
of
(Chapter
be accounted
can
abundance
hours
baboons
assumes
This
Mt. Assirik,
at
food
for
model
This
the
opposite
availability.
food.
the
waking
of
proportion
record
find
the
can be eaten
was high,
increases,
abundance
support
food
that
the
Mt. Assirik
from
any
1974).
by rainfall,
Westoby's
However,
more important
the
to
to
seem
and
Westoby
as food
that
prediction
decreases,
Data
food.
than
more available
time of year (Pulliam
abundance,
more low-quality
ate
was overwhelmingly
as measured
but might
in the dry season,
was abundant
may change,
strongly
assumption
foods
understood
available
the
seasonal
is
to
probably
animals
in
the
Western
in
growing
season
19,73).
If
the
feeding
environmental
concerned
this
record
changes
there
than
is
true
for
with
is
in
the
the increased
at Mt. Assirik,
the
which
wet
season
optimal
itself.
8.21
non-growing
is
foraging
a
season
diversity
reflection
theory
(eg
has
of
not
CHAPTER9: Social
Organisation
INTRODUCTION
Work
9.1: Review of Earlier
The literature
is
baboons
other
on the
1962a,
Hall
1962b,
1969, Saayman 1971a,
1979,
Bert
their
to
returned
Their
forests.
(1969),
the
troops
about
observations
who described
In
to
of
a homogeneous
a report
on a short
100km from
Mt. Assirik,
"some degree
"the
bonds
weak and allow
study
Dunbar
which
the
survey
confirmed
Zoo, Chicago,
adult
that
that
his
his
units
resembles
that
intermediate
those
between
[to
dry
Gaillard
saw evidence
the
males]
are
independence".
lived
foraging
of
in
groups
social
females,
by DeKeyser (1956)
that
this
troop. " They went on
with
P. papio
of
P. hamadryas
9.1
composed of a single
troops.
had a social
and
this
group in Brookfield
associated
(quoted
during
groups
made on a captive
observations
(1973, p186) concluded
on the
that
and
p328)
females
were embedded in multi-male
outlined
(1972,
considerable
(n=72)
male and one or more adult
reproductive
a io
out a 9-week survey of P. papio in Senegal.
sightings
P. papio
P.
of
the GPN de NK, at a site
the P. papio
the
been
subgrouping.
and Nathan
attract
females
Dupuy
of
in
P. papio
within
Boese (1973,1975) carried
He reported
of
in
has
Senegal's
south-east
those
with
troops
and speculated
band without
of substructuring
comment that
relatively
agree
and Stolba
They commented
in
visibility
1967a,
little
in which
sequence
appearance
poor
1966a,
in the wild.
evening.
the
1980,
Rowell
comparison,
the
to
1975, Muý.ller
1980, Sigg
in
be an adaptation
might
1980,
sites
sleeping
anarchic
unstructured,
the
recorded
Kummer 1968a,
1975b,
and Kawai
of P. papio
organisation
(1967b)
et al.
1975,
Hausfater
By
and
1979b, Dunbar and Dunbar
Mori
1976).
the
1965,
1978, Sigg
Seyfarth
Wrangham
on the social
published
for
1971c,
Altmann
1971,
al.
of
1966, DeVore and Hall
Rasmussen
1975, Ransom 1971,
Ohsawa and Kawai
press,
1965,
1963,1965,
Kummer 197ka, Kummer and Kurt
Stolba
1973,1978c,
and DeVore
Hall
et
1959, Crook
1980b, Bolwig
system
and mating
Aldrich-Blake
1963, Dunbar
DeVore and Washburn
organisation
(eg
extensive
1970, Anderson
Altmann
social
young.
This construction
in Boese 1975).
organisation
the
These
savanna
Boese
which was
baboons
(P. anubis, P. cynocephalus,
Boese's
occupy
represented
baboons,
(1)
an
P. papio
in
1970, p261).
between
to
similar
derived,
from
radiating
for
form
a
(or
form
which
1970b for
must
At least
in one-male
baboons
the
savanna
the
which
baboons
to
the
through
passed
the
or those
Altmann
ecological
baboons
these
Harding
of
one-male
problem,
and,
possibly,
those
either
groups
a
ecological
we would
key
its
to
thus
prep,
Dunbar
1973).
in
Boese's
account
In the
leading
presented
solution,
troops
or P. cynocephalus
pressures
be
which
T. gelada,
and van Praet
1970, Western
for
baboons
in multi-male
live
of
floristically
of
1971,1973;
Nagel
of
troops
within
habitats
in
1976, Demment in
explanation
formation
than
1976,1977,
the evolution
for
appearance
exist
to ecological
and Harvey
of some baboons which
and Altmann
a unitary
its
place,
goes beyond baboon
is adapted
explanation
for
taken
interest.
theoretical
Clutton-Brock
(P. hamadryas,
at GilGil,
at Amboseli,
(see
more diverse
groups
has
evolution
organisation
now account
some of
1974c)
(eg P. anubis
for
from
savanna
of Boese's suggestion
is
then
and structurally
search
P. papio
the
and
species
the
be of great
social
it
reviews),
groups
and Dunbar
to
near-parallel
also
If
as we believe
conditions,
live
P. hamadryas
to
also
that
conclude
ancestral
versa),
vice
would
taxonomy and evolution.
richer
the
similar
The second major implication
P. papio.
organisation
or
some remarkable
the reasons
one-male
P. anubis
form.
Alternatively,
Crook
might
and P. ursinus),
between
social
and
either
species
P. papio
we
link
was closely
P. hamadryas
intermediate
Were its
position,
P. papio
(as are P. anubis
similar
evolutionary
such that
Papio
(2)
(Hill
intermediate
an
implications.
be morphologically
seems to
and P. hamadryas
important
two
are morphologically
P. papio
and
has
conclusion
P. hamadryas
P. ursinus).
with
the
the
to
both
a
of
adaptation
P. papio.
However,
example,
exists
whether
(1975)
states
that
Anthoney
some adults
opposite
some doubt
interacted
sex,
the
than
more often
data
showed
that
9.2
while
Boese
expected
with
adults
did
is
(1973)
certain
not
For
correct.
had
shown
adults
form
that
of the
exclusive
relationships
with other
troop
must accommodate and perhaps
like
respects
for
assumption
baboons
shown that
other
this
is
that
be compared
These societies
belong
in
behaved
at Mt. Assirik
The
elsewhere.
social
organisation
in
general
these
of
organisations
of
to:
The hamadryas, in which one-male groups whose memberships are stable
groups
one-male
does not vary
peaceably
at
and Kurt
1963)
The bands
themselves
always
(Stolba
1968a),
but
consisting
1979)
always
The gelada,
in
units
to
a
or
foraging
into
groups
at least
small
other
part or all
form
herds
with members of others
The savanna baboons.
as the troop
(Hall
and whose size
Kummer
of
clans
each
parties
1979).
(Kummer
1968a).
male
one-male
are
groups
stable
to
Groups of one
of a band, or parts
(Crook
not
(Kummer
a unit
foraging
related
but
The band often,
as
1963).
of several
1966, Dunbar and Dunbar
(Ohsawa and Kawai 1975)
(Crook 1966, Ohsawa and Kawai 1975).
changes seasonally
The herds graze cohesively,
1963,
(Kummer 1968a, Stolba
1975), whose membership is somewhat variable
and whose size
more
whose memberships
with
and
bands
(Kummer and Kurt
of which have a common home range.
to
is closed
(Kummer 1968a,
site
one adult
groups
associate
making up either
come together
two
comm).
pers
one-male
1975),
stable
not
fragments
contain
which
(Ohsawa and Kawai
bands,
is
Stolba
other
Kummer and Kurt
1968a,
sleeping
two one-male
of one or
with
associate
troops
made up of
moves
there,
Such parties
male units,
are
1976,
once
form bands all
form
whose membership
(Abegglen
animals
(Kummer
they
where
1980),
These bands may meet other
seasonally.
sleeping-cliffs
1980),
1963, Sigg
(Kummer 1968a) whose membership
in bands
whose size
Muller
3)
the variability
the known social
with
(Kummer 1968a, Kummer and Kurt
2)
the
at
3).
baboons
of
for
account
P. papio
species
chapter
can reasonably
baboon troops.
1)
has so far
thesis
of the troops
organisation
size between wet and dry season (Chapter
This
many
1975 p65k-660).
proposed for the social
Any explanation
Mt. Assirik
(Anthoney
adults
members of each one-male
group mingling
(Crook 1966, but see Mori and Kawai 1975).
The social
unit
baboons
these
can be regarded
of
and DeVore 1965), which usually
does not
normally
9.3
fluctuate
forages
seasonally
cohesively,
(though
see
Anderson
1980a, 1980b).
body
main
occasionally
a
4)
day
the
of
Small
(Mu*ller
(Anderson
more
composition
and do not
Blake
1971, Hall
et al.
suggests
that
one male
groups.
baboons
these
troops
5)
P. hamadryas
by
described
groups
P. anubis
both
in
associating
some of
detach
themselves
1973, Sugawara
contain
troops,
P. papio
of
within
presumably
the
of
whose
which
are
correspond
to
P. hamadryas-like
troop
is
foraging
while
constant,
(1980).
one-male
groups
some of
the
1979).
It
main
is
animals
foraging
frequently
(Nagel
day
or
more
a
for
often
was
one-male
with
and
composition
variable
such
whether
clear
Stable
Separate
troop.
group,
not
organisation
social
Muller
and
other
with
from
an adult
(Aldrich-
male
troops
would
savanna-baboon-like
a
parties,
for
or
unstable
of
organisation
is not known whether
(1979)
Sugawara
associate
1966a)
of P. hamadryas.
that
x
social
The behaviour
exist.
the
and Saayman 1970).
group
it
Rowell
are
an adult
in multimale
The multimale
to resemble
said
live
but
parties
contain
the
of
model
1971,
al.
and Saayman 1970)
Such
necessarily
the band of P. hamadryas,
sleeping
1980b).
from
may separate
et
1980, Stoltz
1963a, Stoltz
(1973,1975)
Boese's
parties
(Aldrich-Blake
troop
overnight
or
foraging
parties
always
male.
9.1.2: Approach
direct,
The
of a troop
organisation
of
the
observer.
and
individuals
When this
troops
whose
DeVore
1965, Rowell
of
such
al.
subject
studies
indirect
of primates
depends
in
a
is
not
are
approaches
to
possible,
not
all
be cautious
troop
much can
and DeVore
their
study
a large
be done
1961),
Hall
interpretation
have been revised;
1979, Anderson
1980b, Kummer and Kurt
and individuals
are
can be used (eg Aldrich-Blake
9.u
not
to
the
habituated
1962b,
the
(thus
Hall
and
authors
some of
see Aldrich-Blake
1963).
recognisable,
et al.
social
proportion
with
although
generalisations
timid,
the
of
recognisable
(eg
recognisable
in
a
upon having
well-habituated
1966a, Washburn
must
1971, Altmann
animals
approach
members are
Washburn and DeVore's
et
preferable,
1971).
When the
various
The
distinction
mating
systems
almost
all
in
this
aspects
In
system.
between
and examine
progression
in
the
the
examine
age-sex
classes
in
data
certain
females,
and
I
Finally
Guinea
for
of
"indicator"
those
baboon
interpreted
This
model
is
site
and predictions
which
social
with
the
in
light
the
the
of
of
in
could
I
social
and
mating
adult
systems.
there.
the
at
the
of
collected
environment
also
nearest
organisation
data
the model which
are made from
are
males
to
relate
moving
chapter
adults
adult
the
consistent
the
identity
between
for
model
Assirik
Mt.
at
next
grooming
of
a tentative
present
In
behaviours
aspects
groups
organisation.
between
groups,
progressions,
troops
distances
inter-individual
or
of all-male
the
mating
foraging
of
movements,
groups,
since
by the
between
social
and
arbitrary,
and transfer
social
of
is
composition
group
organisation
influenced
are
the possibility
interactions
context
the
periods,
future
the
social
chapter
organisation
discuss
composition
neighbours,
following
I
Intertroop
discussed
made between
the
social
chapter
association
have
and
of
this
together.
I
that
field
form the basis
of
research.
9.2: Foraging
Groups: Introduction
When foraging,
(Aldrich-Blake
units
smaller
of baboons of the genus Papio divide
many troops
et al.
1960, Kummer 1968a, Stoltz
into
1971, DeVore and Washburn 1963, Hall
and Saayman 1970, this
1973, Stoltz
and Keith
study).
the
In P. hamadryas
probably
similar,
1968a).
In
by
(1980)
ecological
suggests
smaller
coalitions
foraging
groups
females
and young.
may protect
DeVore
1965,
that
than
Rhine
the
social
such
groups
food
preferred
Furthermore,
against
as
feeding
competition
predation
and Owens 1972).
9.5
For
not.
authors
suggest
by their
of
all
Wrangham
that
females
adult
or
may displace
we may predict
several
Animals
example,
coalitions
sources;
several
may be
parties
large
contain
(Kummer
group
one-male
is
groups
foraging
are
in
and social
of
composition
females
expected
animals
of
the
is
both
pressures,
adult
from
other
basis
baboons
a way that
in
predation,
since
savanna
influenced
the
foraging
of both
composition
that
presence
age-sex
or
adult
more
adult
males
(eg Hall
and
classes
may
therefore
through
seek to forage near adult
the foraging
Some of
groups.
foraging
P. hamadryas
resemble
females,
than
the
baboons
troops
(eg
P. papio
(Boese
1973,
data,
P. hamadryas
and those
This
Assirik
three
males
adult
that
Anderson
Nathan
but
less
is
savanna
1980b)
and
in
there
are
no
Although
in
no
in
noticed
foraging
parties
of
probably
that
foraging
males
are
rare
in
expected
at
random,
in
foraging
groups
adult
common than
likely
are
than
groups
the
composition
other
containing
groups
be
to
in
found
a higher
of
proportion
no
other
possibly
with
than expected, and
likely
females
adult
females,
males and adult
adult
are
containing
Mt.
at
expected,
females
adult
of
propositions:
groups containing
that
1972).
baboons
savanna
young, are found more frequently
3)
al.
difference
investigates
section
foraging
2)
1971,
common, although
and examines
that
et
adult
frequently
more
been
females
adult
rather
have
and
the
occurring
containing
groupings
principal
of
groups
outwardly
baboons.
the savanna
1)
the
containing
but
Such
therefore
should
with
males
Aldrich-Blake
comparative
P. hamadryas
parties,
adult
Dunbar
P. papio
of
foraging
by chance.
expected
groups
groups
and single
young,
males, who would then be overdispersed
in
found
be
to
than
more frequently
groups
in
but
expected
frequently
less
females
adult
foraging
than
expected.
9.2.1: Foraging
Groups: Methods
A foraging
group
spatially
apparently
members were
(i. e. not
over
much of the
because
large
groups
Kummer's "two-male
male
would
field
Groups
impractical.
have
between
moving
resting
or
engaging
area
of
might
teams")
biological
as a group
from
separated
either
food
was defined
in
made a strict
more
than
other
any
food
of
sources,
social
baboons,
most
Poor
activity).
of
spatial
from
excluded
this
be made up of multiples
of one-male
in which
of more than
the
significance
9.6
appearance
which
would
baboons,
five
gathering
or
criterion
5 were
two to
from
of
or
whose
eating
visibility
separation
analysis
groups
(eg
one adult
be misinterpreted.
when a foraging
Whenever possible,
for
frequencies
distribution
multinomial
females.
males and adult
a 3x3 matrix.
each cell
of the matrix
from
were calculated
known proportions
the
age-sex
by whether the
were classified
into
using
its
group was encountered
such records
be tallied
could therefore
Expected
more than one one-male unit.
none, one, or more than one adult
group contained
Results
All
was recorded.
composition
the
to include
groups were less likely
Smaller
each age-sex
of
in the population.
class
Groups: Results
9.2.2: Foraging
The
troops
usually
progression.
occasional
Coordination
of
grunting
by
or
cracking
processing
troops
The
independently,
foraged
itself
split
often
known
were
into
split
kilometres
many
foraging
to
parties
through
the
out.
quiet
of
course
which
subtroops
subtroop
each
apart,
(Sharman,
When
and spread
in
made
with
forage.
been
may have
cohesive
file,
to
parties
noises
a
single
began
small
parties
sometimes
into
as
splitting
or
they
up into
split
in
roughly
out
between
movement
food.
fanning
in
sites
sleeping
moved
troop
the
in woodland
foraging
they
before
bunching,
off
set
Frequently
from
and Hall
Whiten
in
prep).
The area
which
over
and
cohesive
recorded
the
about
parties
There were 635 animals
Their
was recorded.
divided
composition
males were slightly
over-represented
Groups
more
common than
containing
(chi
expected,
adult
squared
at
(Table
and
males
least
= 24.6;
than
one
two
df
= 1;
When foraging
in
areas
group
usually
became
and
Nathan
(1969)
groups whose composition
shown in
is
Adult
9.1.
(Table
9.1).
groups
male
adult
groups
Table
were
contained
more
slightly
no adult
females
foraging
groups
9.2).
adult
adult
hard
a day or two.
for
in foraging
and foraging
than expected
more often
Both
containing
very
Dunbar
in the 215 foraging
age-sex
was normally
the
plateaux,
indistinct.
of P. papio
a troop
that
spread
500m x 250m.
such as short-grass
of good visibility,
more
troop
coherent
was typically
but
to determine,
the
females
males
p<0.001
9.7
found
were
more
in
frequently
and chi
squared
than
= 9.7;
expected;
df
= 1;
Table 9.1:
Crude Age-Sex Distribution
of Animals Sampled
Groups - Expressed
Foraging
as Percentages
are 95% binomial
in parentheses
Figs
limits
confidence
Age-Sex
Class
_I
F;
Y1
9.2:
Number of
Table
IForaging Group
I
{ 30.6 (28-34)
28.7 (26-32)
40.8 (38-44)
122.7
132.1
145.2
I obs
exp
I obs
exp
i
{
85
none
;
84
one
more 1 46
66.8
87.7
60.5
1 77
1 87
1 51
97.3
86.9
30.8
(Table
respectively)
were
males
other
containing:
groups
1 215
also
significantly
so,
were
adult
single
a result,
one other
df = 1;
and
than
often
(chi
squared = 3.21;
adult
male more frequently
one
groups
animal
from
at
another
9.8
least
groups
(Table
one
age-sex
not
and
9.3).
They
statistically
males and a
(Table
As
9.3).
at least
containing
(chi
or
p<0.001
adult
ns)
than expected
to prediction
containing
females,
other
df = 1;
adult
contrary
no adult
but
groups containing
males were found in foraging
p<0.001)
in
respectively)(Table
expected,
foraging
expected
df = 1;
squared = 18.51;
p<0.05
in foraging
female
Foraging
male
more
and either
males,
(chi
than
frequently
more
df = 1;
squared = 5.36;
found
9.3).
adult
female
more than one adult
chi
found
Adult
Numbers of
1 Males
Females
1 215
Adult
(20-26)
(29-35)
(42-48)
Various
Groups Containing
Males and Adult Females
Foraging
I
p<0.01
Population
squared = 24.59;
9.3).
adult
class
female,
were
one
seen
adult
on
45
Table 9.3:
of Three Age-Sex
Groups of
-9-Compositions
Classes
Animals
Number of
{ obs
exp
Foraging
Female { More Ad. Females
Females I One Adult
No Adult
in
found
chi
I obs
41.19
68.77
I obs
chi
exp
75.28'
35.12
10.62
5.68
115
23.22
2.91
50.49
6.77
21.24
9.55
3.21 116
9.04
5.36
1.64
1.76
9.18
2.01
56.85
6.45
I1 22
1 34
One Adult
M1 26
28.83
0.28
1 46
34.86
3.56
Male
FI
46
34.86
3.56 1 32
1 44
38.92
0.66
-
-
-1
32
48.12
More Ad.
M1 57
32.48
18.51 1 34 25.03
Males
FI
Y1
16.04
16
1 15
Y1
Table
29
5.40
-1
10.41
9. ':
Number of Animals of Three
Groups of Various
This
table
represents
Age-Sex
{ No Adult
Class
M
F
Y
obs
exp
1 83
-
61.31
--
1137
121.01
= No Adult
Mi--F1
Y
occasions,
expected
predicted
Females
69
1131
which
(chi
One Adult
lobs
exp
7.67
1 80
184
59.89
87.68
2.11
1 93
118.36
frequent,
While
adult
df = 1;
86.91
85.35
108.28
though
ns).
chi
5.43
Male
0.00
0.36
5.90
not
10.50
0.51
9.3
Table
of
jobs
6.75 1 31
0.15 1 98
1 29
Females
chi
exp
32.26 0.05
134.95 10.12
58.37
14.78
I More Adult
Males
1107
1 35
1 45
66.55 24.59
9.68
20.81
9.23
28.72
significantly
This
males were found
9.9
1 19
I1
Female I More Adult
One Adult
187
19.35 178
5.50 1 83
was less
totals
the marginal
I7
Fo
d in
Table
Classes_foi
sitions:
Males
116.47
160.74
11.63
10.67
Chi
squared = 0.61;
trend.
Age-Sex
chi
f--8.94 147
17.58 121
I--i--Y1 76
No Adult
Males
exp
does not
in
so,
than
support
such groups
with
the
a
frequency
df
= 1;
(chi
ns)
adult
= 0.32;
squared
females
another
which
expected
by
females
were
unlikely
df
= 1;
p<0.05)
than
expected
likely
less
were
that
to
similar
female
adult
was
chance
(chi
squared
be
found
to
present
in
be found
(chi
in
them
because
principally
to
= 0.32;
in
groups
mixed
14.81;
=
squared
adult
df
1;
=
p<0.001).
(1968a)
Kummer
Blake
(1971)
contained
al.
on
Adult
groups
the
for
to
unlikely
than
the
males.
to
troop
was
do so
could
system
be
still
would
was
visibility
males
foraging
composition
of
10),
contain
one
adult
and those
groups
were
rather
social
groups
differed
no adult
free
were probably
Foraging
male.
found
those
males
The
but,
to
in
that
that
more
the
favour
those
of
likely
found
P. hamadryas.
of
foraging
differently
groups,
social
were
in
females.
to associate
instead,
observation
9.10
from
unlike
groups,
in
results
similar
apparently
This
males
adult
no adult
containing
unlike
animals
contexts.
the
might
more.
once
of
over-representation
sight,
occasionally
feed
to
starting
I moved into
males
adult
and may have biassed
groups,
and
two
adult
A mating
where
after
shortly
and
before
baboons
(Chapter
groups
Reserve,
by Aldrich-
male
females
the
with
the
2).
Mt.
attempted
while
how an adult
oestrus
moved off
especially
adult
summary,
the
of
no
males
adult
(Chapter
Gombe Stream
at
while
foraging
savanna
Since
more
a
containing
In
imagine
to
but
females
visibility
normally
accounts
of
sample
was
that
adult
consortship
animals
for
presumably
hard
restricted
females
me
of
at
seen
parties
females
adult
parties
comparable
males,
foraging
of
group.
(eg Ransom 1971).
immature
watch
in
it
as
poor
while
is
temporary
feasible,
in
of
conditions
equally
it
no adult
baboons
of
one-male
foraging
small
improbable
a harem
over
Indeed,
foraging.
it
makes
control
maintain
based
but
females
groups
be a complete
30% of
limits)
observation
under
would
which
foraging,
when
normally
42% (95% confidence
et
This
that
15% and
adult
contained
25% to
shade
between
contrast,
Assirik
Discussion
reported
in
together
sitting
By
Groups:
Foraging
9.2.3:
to
foraging
were
contain
groups
S
than
expected
females
suggestion
foraging
in
by more detailed
females
that
each
other's
likely
than
adult
males,
although
adult
males
more
were
that
but
company,
data.
the
of
imply
to
seemed
examination
containing
groups
female
adult
be found
to
likely
were
supported
not
no
contained
this
There
this
trend
be
found
was
was some
be
to
expected
adult
found
in
was
not
significant.
The
suggests
found
affiliation
between
two adult
defense
groups,
or,
other
adult
of
access
Since
than
frequently
the
male,
that
adjacent
but
In
suggestions
wish
classes
to
stay
to
be parted.
being
found
collection
Altmann
next
of
the
(1979)
in
be classified
males
would
his
"Type
III"
as an adult
individuals
until
in
are
Introduction
Assirik,
do
whom they
certain
age-sex
Methods
is
taken
described
point
fulfilled
progressions,
male,
be
progressions.
a census
censuses
in
result
might
troops
might
from
individuals
in
the
where
individuals
parties,
were
passed
Most
possible,
be found
to
tend
more
one adult
is
It
restrict
found
were
more than
groups;
particular
9.11
I
adult
Between Age-Sex Classes:
data.
mutual
to
Mt.
each other
animals
in
speculative.
at
censuses
together
unsupported.
foraging
to
adult
coalitions
social
that
other
that
formed
containing
Such attraction
in which
The sequence
to
by which
The methods
could
into
(1)
by foraging
coalitions
remain
like
close
Associations
9.3.1:
sexual
at
containing
benefittgd
females.
between Age-Sex Classes:
fragmented
expected
seems
segregated
environment
an
typically
not
such
temporarily
Association
9.3:
in
nor
were
also
adult
groups
social
hypothesis
males
these
recognised
associated
females
in
expected
males
groups
forage
together
females
adult
hypotheses:
females
adult
to
males
adult
second
however,
and that
that
foraging
or might
coalitions
(2)
neither
in
to
Since
two possible
are
predators,
against
these
foraging
tended
males.
adult
expected
there
males,
form
might
males
than
frequently
more
least
in
observation
that
adult
in
female,
in
Chapter
3.
in
the
was preserved
the
criteria
that
every
juvenile,
used
individual
or
infant,
by
its
and
relative
because
of
always
clear,
B in
A followed
been
between
The
ventrally
transitional,
they
were
matrix
(1.03)
frequency
The
to
unlikely
is
error
affect
is
not
and the
that
therefore
the
results
and Fagen and Mankovich
The
frequencies
frequencies
expected
"null",
could
null
never
by
this
significantly;
(1980).
with
other
(black,
M.
and
pregnant
reproductive
follow
status.
treated
were
One cell
null.
in
expected
frequency
expected
it
since
Brown
as
be followed
could
statistically
see
too
seemed
(1974),
Enke
)
in
the
resulting
on
the
basis
9.12
juvenile
infants
as
ignored,
was
infant
an
infant
logically
its
same as
female
pooled
which
and its
empty,
adult
recorded
progressions
end of
female
were
unknown
of
(F),
juveniles,
older
class,
age-sex
introduced
the
females
adult
start
(Note
the
with
adjacent
younger
together
real
were:
carrying
were
A
analysis
(E),
independent
classed
a
of
status
(0),
or
class
age-sex
the
oestrus)
and
followed,
pair
in
and
sequence.
enough,
classes
of
were
pooled
a single
follow.
resulting
(1977)
the
analysis
age-sex
classes
with
pooled
were
or could
(0).
were
if
female
separately
brown)
and
females
all
have
every
juvenile
older
Some
analysed
Thus
categories.
the
be
to
infrequently
exist.
adult
(S),
male
M.
infant
and
(D),
dorsally
subadult
might
be in
to
binoculars.
linear
unknown reproductive
of
not
a complicated
frequently
used
saw a
was
of
a single
within
used
animals
observer
effect
A need not
classes
age-sex
(assumed
skin
infant
(V),
though
adult
sexual
an
this
sequence
female
carrying
For
of
with
positions
B
class
and
the
However,
classes
(M),
to
progression.
age-sex
The
swollen
females
the
was tallied.
male
Subadult
the
that
a progression
sequence
age-sex
class
age-sex
animals
(Y),
B in
width,
lateral
was reduced
resulting
follow
to
association
adult
this
of
was
sample
in
foreshortening
a census
animals
to,
near
found
were
with
of
pattern
shifting
so
relative
the
given
large
metres
paths,
whose
passing
recording
This
analysis.
several
The 80 largest
was known.
progression
parallel
especially
in
the
were
roughly
animals
Furthermore,
even
this
several
stream
If
in
progressions
most
following
in
used
were
censuses
position
of
matrix
the
were
proportion
compared
of
each
with
the
age-sex
Figure 9.1:
((Obs-Exp)/Exp)
Deviation
from expected
in
frequency
with
followed
in
which
given
age-sex
class
was immediately
M (adult
females
progressions'by:
of
males) F (adult
any
J (subadults'and
juveniles)
12 (brown
reproductive
status)
infants)
infants).
and I1 (black
12
0.5
black
infants
juveniles
i-
males.
MFFM
M
12
TFF.
12
O_tT
J
F
12
females
0
ý,
_
1a
ýý
JJ
12
i
brown
infants"
in
class
the
follow
to
classes
The
Adult
(chi
progressions
58% of
them,
in
last
the
of
juvenile,
carrying
infant)
were
and
squared,
1;
=
than
in
summarised
(30%)
classes
on which
either
an adult
male.
This
which
significantly
the
of
than
same age(chi
by chance
alone
for
p<0.05).
which
by adult
unknown
squared
follow
females
expected
of
(chi
4.64;
=
older
juveniles
the
population
in
These
relationships
may
method
used here
the
age-sex
expected
between
than
greater
have
been
was unable
classes
by chance.
9.13
an
were
(30%)
following
association
there
females
oestrus
immediately
below,
the
of
frequency
their
p<0.05).
60% were
Several
frequently
= 1;
141 occasions
the
males
= 1;
are
9.1.
Fig
statistically
discussed
df
to
first
df
than
males
tended
the
= 1;
male,
Females
adult
head
younger
frequently
follow
the
juvenile,
followed
p<0.01).
from
expected
4.13;
=
squared
censuses,
and subadult
p<0.05)
frequently
to
males,
also
more
= 1;
tended
status
reproductive
were
older
would
adult
in
(adult
an animal
they
but
classes
df
follow
than
ventrally
= 7.457;
dorsally,
to
likely
dorsally
infants
squared
Of
all
infants
carrying
carrying
more
for
p<0.01
an infant
of
respectively).
baboons
of
start
df
= 9.61;
squared
was
10%, which
8.84;
=
both
p<0.025
classes
more often
significantly
sex class
not
female
= 1;
df
at
in
rear
squared
(chi
progressions
age-sex
nine
being
under-represented
were
the
at
up the
(chi
expected
5.19;
=
squared
the
adult
Females
position
and chi
Five.
by
pictorially,
progressions
p<0.005)
brought
males
juveniles
in
= 1;
df
than
and ending
over-represented
= 36.08;
subadult
while
Young
p<0.005
(chi
age-sex
Results
starting
were
males
more often
p<0.005).
df
various
was represented
Classes:
animals
squared
was significantly
(chi
classes
Age-Sex
of
class
age-sex
random.
and
age-sex
between
Associations
9.3.2:
of
other
the
of
chi squared as a measure of association.
using
not
The tendency
censuses.
eighty
being
or
the
expected,
seen
these
followed
two
classes
but
for
between
attraction
in
was
reasons
two
the
to detect.
were found
to
These results
follow
are
others
less
summarised
in
Figure 9.2:
in group progressions.
between various
Association
age-sex classes
towards
Arrows point
Each line represents
of chi squared.
one unit
age-sex class that was followed.
For clarity,
any given age-sex
no lines
are shown representing
class following
another member of that age-sex class.
Note, that adult animals tend to follow
and immature
other adults,
(with
tend to follow
the exception
other
of infants)
animals
immatures.
D
V
Fý
Iý
0--%
--S
D female + dorsal infant
V female + ventral infant
S subadult male
E oestrus female
F other adult female 0 older juvenile
M adult male
Y younger juvenile
I independant infant
Fig 9.1.
The data
than
with
(Fig
9.3.3:
et
data
(1971)
al.
among themselves
females
of
Oestrus
females
while
were
followed
other
infants
were also
123 of
affected
the
the
"following"
with
similar,
rarely
and older
infants
thus
travelling
between
with
age-
the
amongst
between
be
same
young
males
the
amongst
on 50% of
in
as
which
and
females.
occasions
(n=40)
Altmann
(1979).
other
However,
their
females
respects
following
females
study
adult
but often
excludes
an
then
be
would
results
are
more than
following
This
The number of
definition
males
defined
mothers.
therefore
the
followed
Altmann
infants.
by Altmann's
males,
his
"following"
adult
in
given
expected.
by other
females
although
members of
not
those
148 "following"
In
similar
associating
being
to
males
was that
than
nominal
following
seems consistent
one of
amongst
on 30%.
mother
infant.
adult
juveniles
Associations
other
Aldrich-Blake
by
adults,
other
and
compared with
females
unknown number of
other
tended
states,
males
seen to be followed
females
associated
qualitatively
association
by adult
frequently
on their
riding
strongest
difference
less
females
each
individuals
be seen with
to
associations
adult
qualitative
with
associating
followed
A
juvenile
and
reproductive
various
The data
The only
turn
presented
P. anubis.
subadult
the
adult
those
with
of
likely
with
The
they
with
less
were
juveniles.
troop
a
and adults
class,
age-sex
more with
who in
animals,
compared
were
for
sex classes
travel
between Age-Sex Classes - Other Studies
was found,
grouping
to
9.2).
Associations
These
tended
adults
and juvenile
subadult
themselves
that
suggest
another
closely
expected,
juvenile,
together.
classes
age-sex
those
of
P. cynocephalus.
9.14
at
in
least
progressions
one troop
at
of
Mt. Assirik
P. anubis
and
9.3.4:
Association
Between Age-Sex Classes:
The associations
between
social
various
reproductive
themselves
this
found
in
but
states
adult
generalisation,
likely
apparently
only
weakly
with
suggest
that
adult
males
need
not
a
harem
females
oestrus
in
males
consistent
P. hamadryas
P. anubis
in
the
the
groups,
P. anubis
females
to
(Mv
males
ler
groups
tend
follow
the
to
to
follow
mate
only
with
to
access
one
interpretation
of
groups,
that
so
more than
females,
more
Nagel
the
which
the
By contrast,
oestrus
1980,
might
male,
females
females
oestrus
adult
females,
1971).
to
other
P. hamadryas
oestrus
Nagel
(Mu 00
ller
males
this
In
amongst
observation
oestrus
society.
1980,
for
compete
for
of
an exception
with
This
males.
follow
bonds
were
little
compete
do not
groups
male
objections
several
adult
constrained
are
follows
female
with
females
clearly
females
adult
maintain
associating
and
be
that
to
Oestrus
males.
distinguish
not
suggested
were
females
would
did
progressions
organisations,
and with
Discussion
and males
frequently
1971).
data
in
than
There
collected
are
Mt.
at
Assirik.
1)
(1980)
Muller
females
anoestrus
is
apparently
the strong
has
P. anubis
followed
characteristic,
and P. ursinus
a
females
followed
males
the males.
are said
Thus
in Mvller's
although
similar
This
1979).
females
oestrus
demonstrated
one study
trend
pronounced
to lead
their
male
(Saayman 1970).
here
classed
swollen
sexual
skin.
This
sexual
cycle,
of
attractive
the
to
mature
1972).
Female
followed
males
males
(Rasmussen
There
is
oestrus
females
males to follow
Females
phase
the
have been a local
P. cynocephalus
consorts
3)
for
show that
savanna baboons (eg Altmann
for
unusual
(Rasmussen 1980),
2)
to
able
more than
tendency
baboons might
of
was
far
in
females
some
of
(Collins
males
more
were
included
P. cynocephalus
evidence
females.
"oestrus"
as
frequently
at
any
all
whom would
pers
with
fact
comm,
than
they
stages
of
and
swollen
were
with
have'
not
Tayler
partially
female
this
been
Saayman
sex
followed
skin
by the
1980).
that
These
adult
pairs,
9.15
baboons
in
formed
which
the
consortships
female
with
was probably
some
at
her
most attractive
be
found
4)
the
on
the
generally
males
in
these
pairs
to
then
censuses.
Finally,
the
progressions
following
2m behind
The observation
been
likely
these
censuses
(see
female
60% of
an
were
to
adult
since
recorded
tended
recorded
Thus
one side
of
a
to
male
as following
the
pair.
females
oestrus
thus
male
of
and
be recorded
might
observed
adult
seen
to
was
females,
Methods).
metres
all
be
to
this
records
receptive
less
tended
If
obs).
against
sexually
was several
with
association
with
as oestrus
that
(pers
troop
laterally
which
animal
and Saayman 1972),
was a bias
by which
method
compact
close
there
have
would
the
of
association
close
(Tayler
male
periphery
case
during
a third
the
becomes
were
in
biologically
important.
Introduction
Runs of Males:
9.3.5:
Progression
order
(Altmann
baboons
savanna
al.
to
not
distinguish
P. hamadryas
in
found
organisation
social
appears
and
1973a, 1979, Altmann
1967b, Boese 1973, DeVore 1965a, Hall
between the
clearly
found
that
and Altmann
among the
1970, Bert
and DeVore 1965, Harding
et
1977,
Kummer 1968a, Popp 1978, Ransom 1971, Rhine 1975, Rhine and Owens 1972,
Rowell
1969, Stoltz
Progression
Papio
takes
adult
male
form
all-male
are
usually
has
a
"runs"
they
of
would
over
population
groups
found
similar
adult
does serve
order
T. gelada,
from
male
one
and Saayman 1970,
at
social
males
however.
a group
from
which,
the
mating
females,
excludes
breeding
group.
These
to
according
head of
should
the
T. gelada,
baboons of
related
of
system,
,
distinguish
to
In
the
Washburn and DeVore 1961).
the
the
with
occur
by chance.
9.16
herd
in
Crook
in
the
system,
excluded
progressions
the
males
(1968),
If
progressions.
of
where
some of
and Aldrich-Blake
existence
genus
all-male
more frequently
P. papio
groups,
than
9.3.6:
Runs of Adult
Runs of
adult
males
there
would
be
where
number
In
of
to
1-p = 0.77,
first
the
reaching
then
which
males
associate
6
(eg
or
Of
(66%)
were
The
sequence
(chi
was not
was due
Thus
recorded.
not
adult
adult
males,
df
followed
the
probability
If
in
male
j
of
is
progression
the
each
p=0.23,
chance
is
position
adult
of
males
before
If
adult
of
large
)
1977).
frequencies
observed
the
exceed
will
7
the
in
by
of
expected
largest
animal
an
than
more
in
5 were
not
than
to
the
fit
value
198
censuses,
which
was
adult
males
not
another
most
pairs
males
expected
males.
9.17
were
were
but
seen
seen
found
(Fig
and threes
expected,
this
being
expected
with
in
the
of
lack
in
of
and
alone
9.3).
one run
found
distribution
geometric
By inspection,
than
males
found
although
of
p<0.001).
fewer
expected
numbers
good
= 2;
rather
longer
males
the
recorded
a statistically
more than
rather
the
(Pollard
males
distribution
= 22.2;
to
then
Results
males
and
at
9.5).
frequency
squared
fit
adult
preceded
themselves
distribution.
geometric
adult
(Table
adult
Males:
assort
(j=0.1.2....
0.77
groups,
of
males
counting
distribution
all-male
by
is
animal
groups
299
the
male
adult
in
was recorded.
introduced
population.
of
analysis.
error
an adult
probability
geometric
Runs of Adult
9.3.7:
not
this
the
in
the
= Piq = 0.23'x
from. the
calculated
the
the
more)
is
which
other
P(J)
defines
in
in
used
largest
seven
another
males,
any position
males
an animal
encountering
q=
adult
of
proportion
adult
in
if
that
The
troops,
one
ignore
may
so
other
to
larger
the
males.
were
adjacent
we
male
adult
records,
seen
in
particularly
excluded
sampling,
an adult
encountering
occur
progressions
relative
random
the
more
replacement
assuming
should
males
adult
large
very
Methods
1853 age-sex
totalling
censuses,
The
Males:
Runs of
6 males
animals
company
that
of
was
were
other
9.5:
Freauen
Table
Distribution
of
Found in
Numbers of
Progressions
Frequency
obs
exp
Number
in run
chi
4.51
66
52.95 I
3.22
3
25
12.18
4
9
198
2
230.23
f
The
in
found
they
would
Adult
i0
I1
10
data
from
the
vicinity
Males:
1
i
1
0.64
0.15
0.03
11 22.22
by chance
large
socially
peripheral
model
in
a multimale
attraction
to
general
were
the
to
seen
pattern
similar
together,
towards
P. cynocephalus
sampling
could
Adult
presented
by the
individuals),
progression
baboons.
seen
predicted
no
contained
to
than
small
group
by
at
in
the
(19651,
Mt. Assirik
and
In
the
distribution.
in
the
proportion
contained
more
not be assumed.
than
9.18
of
8 adult
because
adult
males
males,
may be
males
hierarchy"
a qualitatively
at random would have been found
geometric
a
showed an
"central
was found
(1979).
with
with
which
males
attraction
adult
a population
between
of
than
of
seem consistent
adult
DeVore
by Altmann
suggest
but
males
blocks
be
to
more frequently
expected
groups,
closely
adult
be
would
This
that
do not
all-male
associate
males distributed
(484
as
data
tend
males
adult
rather
males
other.
each
savanna
adult
The
alone.
travelling
that
suggested
other
males
which
Discussion
Mt. Assirik
of
13.49
2.80 I (13.73)
1 299
Runs of
Male Baboons
sq
i
1
5
6
>6
9.3.8:
Adult
is
the
latter
data
on
example
in shorter
runs
the
sample
low
(0.15)
so that
is
and
replacement
Intertroop
Encounters:
Introduction
When
troops
baboons
9.4:
two
between
interaction
1976),
while
fights,
(1976)
had
tended
to
(Ransom
peaceful
Hall
a few
report
for
and,
herds
(Rowell
(Hamilton
Crook
1976,
al.
were
1966),
pers
comm),
(Saayman
mingled
even
frequently
(Aldrich-Blake
Rasmussen
or
studying
intertroop
1973),
peaceful
troops
which
separated
et al.
between
Males
that
to
establish
Saayman 1971a,
1971c,
exchanged
members
to
transfer
relationships
with
are
about
If
normally
occur
presumably
when I
that
at
the
they
involved
have
(Aldrich-Blake
splitting
back
the
see
were
front
two
their
of
out
of
members
interactions
prep,
to
troop
the
near
could
of
troops
peaceable
troops
natal
other
are
troops
during
transfer
peaceable,
Rasmussen pers
may
comm).
Methods
difficult
of
majority
moving
in
Encounters:
was extremely
the
these
(Manzilillo
Intertroop
process
resources
highly
while
as
1966a).
be easier
may well
defensible
the
that
interpreted
been
and
1971c),
imply
to
the
interactions.
intertroop
restricted
over
are
have
troops
in
or
1971, Rowell
It
T. gelada,
have been taken
recently
9.4.1:
in
encounters
et
interactions
certain
of
encounters
males
and Harding
describe
Kummer 1968a,
1963a,
they
(Paterson
invariably
or
interactions
competitive
even
1971),
(1973)
that
workers
peaceful
adult
1966a).
Aggressive
start
seasonally
1962b,
1971,
al.
while
were
which
encounters
Saayman
Cheney and Seyfarth
Nagel
troops
Other
troops.
(eg
(1965),
the
which
al.
overt
1974,
al.
in
encounters
et
involve
et
troop
resulting
(Hamilton
territorial,
other
that
the
other,
(Buskirk
males
the
believe
other
avoid
territorial
DeVore and Hall
to
reason
be
observed
away from
Saayman 1971a).
1977,
each
necessarily
adult
have
meet
may
not
between
females
adult
troops
though
others,
Many workers
herded
(a)
the
especially
1971c).
et
of
was normally
the
of
or
groups
two
observe
9.19
out
troop,
leading
moving
different
intertroop
of
and
sight,
most
flanks
closer
troops
of
(b)
and
encounters
the
together
and
since
encounters,
not
troop,
I
was
subtroops
I was
would
(c)
and
rarely
of
a
larger
troop.
This
generally
an indication
in
obs)
the
which
included
observations
largest
intertroop
have
been recorded
would
report
when the
encounters
vocalisation,
this
were
were made.
(250)
number
including
troops
in
to have happened
the
exceeded
when the
season,
therefore
were judged
behaviour,
(pers
dry
and I
Aggressive
troop.
any
the
in which
season
a progression
by characteristic
baboons
3),
interactions
in
animals
occur
the
of
Peaceable
of
(Chapter
smaller
in
true
was particularly
are
in
number
known
to
accompanied
other
species
of
at
Mt. Assirik
had
of
apparently
stable
it
occurred.
Intertroop
9.4.2:
In
the
Encounters:
membership
(Chapter
apparently
peaceable,
the
into
4).
One
sleeping
site,
and
sleeping
site
(Chapter
occurred
near
sleeping
sites
such
or
on
In
the
smaller
troops
of
membership
accompanied
The
(described
the
season,
troops
these
in
in
were
were
either
was
loud
barking
et
with
1974,
al.
seen
at
9.20
that
nearest
to
the
encounters
observed
to
share
from
the
other
is
not
All
though
in
Hamilton
in
known whether
the
such
fissions
and
prefaced
and
and pers
intertroop
aggressive
These
the
other
usually
press
4).
and
each
constant.
Mt. Assirik
season.
It
(Byrne
the
(Chapter
rejoined
peaceable,
of
membership.
fragmented
fragmented.
associated
not
troops
moving
together
separation
original
frequently
later
Buskirk
its
large
fragments
behaviours
After
from
similar
were
were
troop
members
moved
other
Troops
sites.
seemed to keep
3
troops
one
km
group
least
occasions.
by increased
Saayman 1971c)
encounters
many
troops
of
At
apparently
larger
resulting
fusions
dry
3).
of
with
2
about
large
resulting
sleeping
at
each troop
troop,
the
associating
occurred
large
these
some members
peacefully
encounter
and.
between
least
at
and
troop.
large
were
Encounters
with
troop
other
troops
season
wet
Results
encounters
1975b,
et
al.
the
context
obs).
of
1976,
intertroop
and
9.4.3:
origin
the
ecology
of
proportion
had
few
for
home range
sites,
(median
length
The
from
protection
interactions
peaceful
of
advantage
home
help
might
range
reason
for
such
easier
for
individuals
9.5:
Between Troops:
Transfer
the
between
transfer
species,
of
is
and
the
between
since
each
gives
roosting
these
there
might
circumstances
be
the
to
in
their
Whatever
relationship.
they
the
made it
probably
troops.
Introduction
the
environment,
troops
probably
5).
elsewhere
interactions,
transfer
(Chapter
1976b)
interactions
the
whose median
probably
would
the
intertroop
to
Under
site.
one
dispersed
low,
communal
at
sleeping
so that
troops
and Vehrencamp
sites
reinforce
if
of
least
at
6),
was probably
and peaceable
peaceful
Irrespective
can
sleeping
if
was a journey
of
8 known
widely
km; Chapter
but
data
more
or
with
the
best
The
were
the
of
baboons,
calculation,
sites
and
the
rest
of
one
high
a
included
share
site
6),
to
with
mean day range
sharing
troops,
both
shared
1.5
=
(Chapter
at
which
would
sites
in
benefit
been a net
have
troop,
Sleeping
(eg Bradbury
predators
water.
sleeping
many trees
contained
site
permanent
a sleeping
the
most
sharing
of
cost
biomass
it
site
the
of
most
a large
troop
ten.
contained
site
By a rough
each
from
20% of
was roughly
of
which
nearest
displaced
being
of
cost
to next
distance
to
5).
in
one night
on about
other
7 of
random,
at
relative
Fable's
from
(Chapter
troops
used
were
sites
least
at
The
sources
come
area
4 other
least
or
may owe their
Mt. Assirik
at
area.
supporting
of
sites
sleeping
sleeping
habitat
capable
troops
the
of
productive
apparently
park,
between
interactions
Peaceable
to
Discussion
Encounters:
Intertroop
baboons
of
varies
which
greatly
by the
influenced
partly
with
ease
males
adult
baboon
across
mating
of
structure
species.
At
herds
least
(Ohsawa
P. hamadryas
Some evidence
between
some one-male
bands
and Kawai
one-male
that
in
group
groups
1975),
or
entire
P. hamadryas
in
but
solitary
one-male
is
there
adult
groups
was provided
9.21
transfer
T. gelada
no
male
do
documented
moving
not
between
readily
between
normally
by experiments
case
of
a
bands.
transfer
(Kummer 1968a)
in
two
which
sleeping
while
male,
its
and
males
troop
(of
herds
may stem
from
units
T. gelada
all-male
succeeding
harem
previous
one-male
entire
a benefit
on,
harems
adolescents,
fighting
to
therefore
introduction
or
adolescence
(Kummer 1968a,
the
well
male
baboons
(eg
Packer
probably
(Packer
easy
1979,
in
p49).
or
after
The
late
in
one
to
a threat
present
(Rowell
1966a,
Tayler
any
female
are
highest
in
agonistic
1980),
but
not
always
likely
to
not
succeed
maximum
achieve
between
all
vigorously
resisted
to
rank
(eg
troops
least
copulating
agonistic
(eg
often
females
Although
rank
newly
(Packer
1975b,
in
the
Manzilillo
later
mating
by
males
who
Rasmussen
more
prep),
period
transferred
1979,
move
troop
Resident
Strum
at
their
in
Transfer
baboons
Hausfater
1972a,
1971a,
own
these
male.
single
with
9.22
In
once
may sometimes
their
into
all
and nearly
1973a).
males
adult
1972).
are
at
Hausfater
back
Saayman
fertility.
high
1979)
prep,
any
males
is
troops
between
Rasmussen
that
Saayman
restricted
in
in
shifting
and
of
movement
baboons
savanna
between
rapidly
1968a,
apparently
1975,1977b,
Manzilillo
temporarily,
is
the
transfer
troops
probable
and is
band,
units,
especially
male,
therefore
must
baboons
savanna
documented
may be so
a new adult
of
one-male
(Kummer
form
males
juveniles
as
other
an
of
and may confer
P. hamadryas
usually
"owner"
transfer
to,
the
from
females
no threat
from
and live
occasionally
The
By contrast,
Many
systems.
of
1975).
can
P. hamadryas
herds,
family
a
units
p112).
Among the
lifetime
of
as adults
adulthood,
in
by the
population
the
of
females,
previous
by
mating
breeding
the
Dunbar
herd.
band
their
represents
occasionally
males
resident
harem
single
early
in
from
and
recipient
the
by resident
T. gelada
which
experienced
periphery
a
group
the
with
the
over
(Dunbar
but
resident
fighting,
severe
with
difficulty
the
on
kidnapping
by
ease
excluded
male
its
strange
the
of
were kidnapped
after
differences
are
taking
in
off,
the
the
the
groups
among a
or
350 animals).
and
males
adult
near
unit
other
driven
between
between
switch
of
adult
difference
The
the
of
male
sleeping
entire
in
females
the
released
was chased away by several
One unit
troop.
were
units
one-male
their
of
males
in
may
prep),
are
they
have
been
those
are
all
all
register
of
recognised
of
other
known individuals
of
known individuals
Transfer
On 8/8/78
individuals,
individual
with
seen
later
was
in
females
male.
9.5.3:
two troops,
this
group.
seen
on
Fable's
Transfer
to
one troop
notice
the
group,
Between
giving
it
its
record
of
this
and
occasions
of
Troops:
characteristic
male
was
prep).
The
and in
this
way a
Any movement
was recorded.
another
(Chapter
immediately
had
to
to
and accepted
3).
been
was
but
before,
seen
be groomed
and
An
tail
recognisable
not
groom
near
450-500
them about
2km together
sitting
by adult
known adult
a particular
by the troop.
Discussion
a previously
on previous
and
was compiled.
to
which
recording
(in
Sharman
recorded,
between
individual
while
periods,
Results
and moved about
have come about
him
given
from
He appeared unstressed
might
age,
was made of
recognising,
in
each troop
social
A register
for
used
containing
several
The observation
tail
similar
any characteristics
date.
in
This
during
for
was also
a characteristic
with
detail
are
Between Troops:
coalesced
male
adult
the
by allowing
lose
to
most
especially
a later
The methods
any recognised
9.5.2:
but
in
them at
individuals
indexing
presence
the
of
costs,
that
shown
those
are
severe
Methods
were scanned
individuals.
such
new males
have
day,
the
identify
to
serve
might
during
times
baboons
visible
all
the
resist
has
prep).
immediate
little
of
(1979)
Packer
in
Strum
and may incur
males,
until
copulations
1979,
therefore
resident
Troops:
Between
Transfer
At
is
presumably,
which,
(Packer
a year
injury.
to
fertile
in.
new males
9.5.1:
about
the
to
likely
most
residents
few potentially
males
debilitating
as
which
for
incoming
benefit
reproductive
in
succeed
resident
to
Resistance
such
to
likely
they
by (a)
unknown
a failure
occasions,
shape.
made all
9.23
or
male
with
a characteristic
the
part
of
of
(b)
other
injury
a fresh
However,
animals
by
the
the
in
time
both
to
observers
his
the
Fable's
tail
first
and
troops
Anecdote's
been
with
the
repeatedly
for
improbable
that
he
possibility
of
a fresh
he
but
being
apparently
recent
if
were
as
they
The
to
serves
P. papio
for
difficult
Awash.
By implication,
groups
are more likely
T. gelada,
to
they
than
lick
between
if
be founded
are
it
with
that
this
on groups
Mt. Assirik
at
this
single
is
example
troops
was
as
P. hamadryas
for
in
of
into
each other.
was
mates
to
or
baboons
between
kidnapped,
on
tail
suggests
However,
P. papio
ruled
them and behave
touch
troops
transfer
is
The
be
his
Other
joined
as
it
and
and had transferred
temporarily
that
seen
sixteen.
to
evidence
troop
was
cannot
to
or
the
Assirik
then,
first
attention
incident.
Mt.
at
the
had
he
study,
occasions.
another
hypothesis
the
reject
the
of
coincidental,
any
transfer
one
been
for
summary,
two troops
on
resting
pay
to
male
had
months
several
In
for
seen,
were seen to
belonged
evidence
anecdotal,
to
on
painful.
when the
troop
Fable's
he
however
seen
injuries
characteristics
missed
injury,
groomed
had previously
male
three
been
had
distinguished
once
remaining
was never
it
prevent
easily
Furthermore,
recorded.
out,
such
the
groups,
one-male
females,
related
females
coerced
at
as in
as
in
will
it
P. hamadryas.
SUMMARY
The
be
individuals
until
may form
in
so that
easily
into
at
their
T. gelada.
is
of
any
P. papio
of
it
social
many complex
the
Mt.
known
social
About
organisations
resembled
organisation.
one in
five
9.24
of
a
of
time,
over
(Dunbar
one
1979; Altmann
on the demography of
observed
(Hinde
that
nor
populations
one troop
as those
surprising
social
Assirik
within
within
ways
clear,
yet
organisations
as large
not
not
depends partly
apparently
perhaps
is
However,
and indeed
groups
relationships
1976),
unlike
This
Members of
troop.
aspects
baboons,
1979).
and Altmann
this
known.
a range of different
can find
P. papio
of
are
of
species
single
the
organisation
social
in
this
study
and Stevenson-Hinde
P. papio
of
does
baboons.
the
savanna
baboons
They
foraged
in
these
foraging
small
groups
not
fit
Despite
in
many
groups,
contained
but
females
adult
no
between
Associations
males,
adult
age-sex
classes
baboons.
savanna
be found
in
were
found
males
were
in
runs
dispersed
they
apparently
was
recorded,
characteristic
were
a moving
lengths
of
randomly
resemble
in
of
which
some
that
through
savanna
the
were
close
the
baboons.
animal
savanna
P. hamadryas.
9.25
was
baboon
to
similar
of
(Kummer 1968a),
clan
in
unlikely
The males
pairs
in
is
which
to
P. hamadryas.
those
found
P. hamadryas
but
that
In
One transfer
populations
tend
if
this
between
easily.
but
to
P. papio
expected
progressions.
accepted
in
males
in
the
also
troops
This
is
not
of
CHAPTER10: Mating Systems
INTRODUCTION
10.1: Review of Earlier
The
detailed
depends
upon its
whether
or not
the presence
Hausfater
strategy
mating
age, its
it
specific
(MuJller
the mating
of P. papio
Our current
view
the
of
Boese's
(1973,1975)
published
Boese's
field
was not
the age-sex
record
any
work
account
of
should
1967a),
we should
breeding
be treated
hesitate
has
male
(reviewed
designed
to
mating
9).
Since
1973),
and since
of
animals
1963,
and Kurt
or
to
in
Rowell
in which
system
one
over
on
but
a mating
rights
largely
structure,
(Boese
(Kummer
find
baboons.
rests
mating
to P. papio
to ascribe
exclusive
are species
on observations
caution
1979a,
systems
in Chapter
parties
based
troop,
strong
of other
briefly
of foraging
of the
variability,
P. papio
examine
troop,
We may therefore
from that
work
with
this
of
the
(Dunbar
on
so
and
1979).
system
mating
organisation
social
captivity
each
composition
differs
probably
in
structure
of baboon mating
1971, Sugawara
troop
a
animals
troops,
Despite
the fundamentals
system
of
other
the age-sex
1980).
1980, Nagel
member
number of
of neighbouring
Rasmussen
that
exists
any
in the troop,
has kin
1975b,
of
the
rank,
and structure
evidence
that
Work
more
adult
females.
In this
Assirik
exist
1)
is
I assume that
chapter
similar
in baboons.
The mating
to one or other
These are,
the
of
five
particular
mating
mating system includes
males.
adult
1968a, Kummer and Kurt
female
bite
or
slap
strays
her
to
1963)
females
1963).
characteristic
females by adult
adult
and particular
from the adult
the
time
males
attention
(Kummer
males
spatial
proximity
by the females, but when an
he
will
male
chase and sometimes
(Kummer 1968a, Kummer and Kurt
10.1
such a
of
and near-continual
Most of
(1963)
females may only mate
of anoestrus
between members of the group is maintained
adult
thought
by Kummer and Kurt
adult
Behaviour
the herding
(Kummer 1968a, Kummer and Kurt
between particular
systems
at Mt.
in brief:
in which particular
adult
the baboons
of
system
system of P. hamadryas, described
(1968a),
Kummer
by
and
with
the mating
1963).
Oestrus
females
will
for
openly
interact
rarely
to
copulation
(Kummer and Kurt
1963).
more than
(Abegglen
hierarchy
males are never seen to compete for
the [sexual]
would follow
(Kummer
1968a).
succession
the adult
"at
females:
males
but
most, a male
female with his eyes"
different
of
(Abegglen
and mingle
that
of a passing
members
in
male
present
1976, Kummer 1968a),
oestrus
swellings
Female
sometimes interact
one adult
There is some suggestion
a dominance
recognise
(Kummer 1968a) and never
with
groups
one-male
1976, Sigg 1980 p276, Stolba
1979).
2)
The mating
are
made up
females
the
grooming
interacting
Adult
with
females
rarely
male
(Dunbar
females
in
with
These
always
associated
with
to
his
of
which
the
all-male
the
with
other
prevent
than
more
one
(Mori
1975).
adult
in
male
with the
any interaction
and Kawai
from
groups,
1979b, Dunbar and Dunbar
interrupts
females
time
differs
T. gelada
to
the
social
a secondary male is sometimes associated
male usually
by
maintained
attempts
male
belonging
1979b).
most
1978a),
P. hamadryas,
unlike
largely
spends
the
on
(Dunbar
is
is
adult
interact
The primary
secondary
those
adult
although
succession,
group.
The
particularly
males,
adult
male
(Dunbar
females
adult
females
cohesion
adult
P. hamadryas.
from
again
the
but
females,
adult
Group
groups
all-male
which,
male which
an adult
1979b).
with
1966)
related
of
with
mate only
(Dunbar
them
groups
of
(Crook
herds
mixed-sex
of
periphery
T. gelada,
in
found
system
of
In
1975).
the
this
P. hamadryas and T. gelada are similar.
3)
In these baboons, groups of
The mating system of the savanna baboons.
related
females,
females
from other
within
matrilines
and Ransom 1971, Strum in prep).
are
frequently
(Collins
the
troop
Consortships,
seen between oestrus
in prep, Hall
amongst themselves
interact
or matrilines,
adult
and adult
males may form special
the female's
oestrus
periods
Rasmussen 1980, Seyfarth
associations
(Altmann
1978).
10.2
(Cheney 1977, Ransom
or mating associations,
females
and DeVore 1965, Hausfater
Ransom 1971, Rasmussen 1980, Saayman 1971a).
and with
males
and adult
1975a, Paterson
Individual
adult
which persist
1973,
females
outside
1980, Packer 1977b, Ransom 1971,
Herding of anoestrus
females by adult
males is rarely
1977) or during
Seyfarth
them in one direction"
interact
frequently
The model
in
baboons.
copulation
females
to
than
again,
(Boese
1973),
5)
so that
males
for
(Boese
but
male
savanna
adult
(Boese
for
present
or
(Boese
1973),
male hierarchy
(presumably)
females
punish
the
succession
An adult
P. hamadryas,
to adult
access
in
and
P. papio
or
with
embedded
in which
T. gelada
not
are
herd
1975),
interact
competition
although
ignore
frequently
"consistently"
or T. gelada.
unlike
to more than one
groups
one-male
females
one adult
P. hamadryas
resources,
between
more
may
1978).
males
never
females
adult
1978) who will
which
P. hamadryas
Oestrus
like
in
Adult
resemble
members and chasing
copulation
and adolescent
adult
troop
for
or present
(Cheney and
encounters
Oestrus
(1973),
troops.
then
would
[of]
in prep, Seyfarth
Boese
due to
multi-male
anoestrus
"gathering
(Saayman 1970, Seyfarth
her invitation
ü)
intertroop
(Saayman 1971a).
with
male (Collins
adult
during
seen except
exists
males
interact
over
never
takes
place
1973).
P.
hamadryas
P.
in
found
anubis
x
The mating system
by Sugawara
troops
(1980).
In
these complicated
societies
by
Muller
and
%wor
behave like P. hamadryas and some like P. anubis.
individuals
some
(1979)
behaviours
of both mating systems are seen.
characteristic
I have chosen to analyse
five
of social
aspects
life,
in
animals
different
compositions
age-sex
of
will
class
social
from those of animals
the
of
adult
(Nagel
other
it
is
animals
depends on both
1971).
Some behaviours
male or the female
that
(Nagel 1971), and finally
on whether
is
for
groups
The
females,
adult
pair
distances
between
and on the
species
the maintenance
of grooming relationships
1980, Saayman 19716).
10.3
one.
depend upon whether
for
have
will
males who are the focus of
the
sex of
responsible
kin
example,
Inter-individual
of adults
the nature
or not matrilineal
Kummer 1968a, Muj ller
the
society
in a more liberal
they or adult
females.
adult
of,
neighbours
nearest
depend upon whether
attention
a harem
in
behaviour
each of which
Small groups of
helps in some way to decide between these mating systems.
involved
Thus
exist
it
is
the
of proximity
will
depend
(Dunbar and Dunbar 1975,
10.2: Social
Groups: Introduction
The composition
to
the
mating
groups of baboons is apparently
of social
in
structure
the
troop.
Nagel (1971)
baboons of different
species,
troop,
where females
were socially
social
groups containing
in
contrast,
to
the
interact
also
10.2.1: Social
[that]
p51)
in
each of
the
the
smallest
neighbouring
to
the
clumps
of
contact
small
radius,
one or
behaviourally
reason
they
might
distribution,
population
than
frequencies
the
with
and
the
10.2.2: Social
Groups: Results
cliffs,
the
of
4m from
were
animals
neighbouring
their
well-defined
in
the
in
clump
was apparently
For this
group
was at
baboons,
even
though
neighbours.
from
calculated
numbers
possible
the
that
implying
used at
in
animals.
neighbouring
2
In practice,
party.
Each clump
group".
from
observed
of their
areas, either
from
known frequencies
irrespective
After
each
"social
was not always
and interacted
neighbours.
isolated
have been less
it
was
between
was also
criterion
was
animal
distance
the detached
sat
with
of... detachment...
greatest
This
detached
of
composition
and another
meant that
distinct
term
and
the
[party]".
its
of
and spatially
Expected
than
frequently
often
behaviourally
time
containing
= 3.37, p=0.001).
[party]
the
as 4m around
more
I have used the
the
larger
the
a distance
Mt. Assirik
with
were free
groups
social
By
male.
females
adult
The criterion
between
poor visibility
at
baboons
49% of
size
groups.
resting
within
although
see as great
the
"recorded
distance
animals
Mt. Assirik,
where
male (t
an adult
4m and 4 times
than
greater
only
an adult
contained
Methods
(1971,
parties
individuals,
contained
Groups:
Nagel
baboons,
of
male, 80% of
by an adult
also
troops
in the P. hamadryas
found that
of P. anubis
three
of
a study
restricted
females
adult
other
with
females
adult
troop
In
sensitive
of
each age-sex
groups
of
the
of
multinomial
in
class
the
sizes,
various
composition.
the baboons left
in the valleys
their
sleeping
trees,
they collected
in clear
amongst the rocks at the base of the laterite
or in areas of bare laterite
10.4
or short
grass
on the plateau
edges
bordering
The time spent here varied
the valleys.
(Chapter
hours
several
in
associated
continual
groups
There
groups
the
many of
more individuals.
within
between
and
if
small,
especially
by
expected
the
no
adult
number
(151.33)
Adult
animals
There
was
so
that
might stay constant
for
groups
large
with
df = 1;
with
more
or
groups,
change
c 1.13;
(136)
(chi
alone
squared
= 1;
df
different
from
the
= 1.55; df
= 1;
ns).
females
adult
(chi
10.3) so that
the number of adult
adult
females was smaller
other
groups containing
squared = 4.33; df = 1;
females
adult
tended not to be found in groups containing
one or more other
10.2).
ns)(Table
containing
of
from
was indistinguishable
significantly
not
Table
The frequency
analysis.
females
groups
in
shown
are
classes
in the
squared
social
was
(Table
p<0.01)
in social
of
age-sex
included
(chi
by chance
females
male together
one,
chance
males
whose
groups
none,
containing
expected
Table
might,
1372 animals,
were
Furthermore,
(chi
more or
time
10 or
group, especially
the 441 social
10.1, in
but
this
to
times in a few minutes.
several
that
from 2 to
of
membership of a single
or
During
movement both
sporadic
half-an-hour
u).
from a few minutes
number
one adult
squared = 7.14;
found
females
than expected
p<0.01).
10.1:
Crude Age-Sex Distribution
of
Groups - Expressed
Social
in
Figures
parentheses
Age-Sex
Class
M
F
Y
Animals Sampled
as Percentages
95% binomial
are
confidence
in
limits
I
Social
i
124.2
Group
i
I
Population
1 31.2 (28-34)
1 22.7
132.1
1 44.6 (42-48)
1 45.2 (42-48)
(21-27)
(20-26)
(29-35)
KEY: M= Adult and Young Adult Male
F= Adult Female
Y= Subadult and other Immature Animals
Social
adult
males
squared,
groups
nor
p>0.05
containing
adult
more than one adult
females
more
in both cases)(Table
10.5
frequently
10.3).
male included
than
neither
expected
In particular,
(chi
no social
10.2:
Table
er of
Expected
Groups
Social
of Adult Males
frequencies
Containing
and Adult
I
Females
137 136.5
none
one 1 187 179.1
more 1 115 123.5
10.3:
Number of
Table
Expected
Animals
1441
from
calculated
exp
'lobs
chi
MI
FI
Yf 153
One Male MI 45
FI
-
f1
65
11.74 1111
116.09
58.87
98.27
3.27 f 97
197
1108
4.60
YI
77
More than MI
one Male FI
YI
81
66.39
3.22
32
24.59
2.23
I
groups of more than 3 animals
the
Comparing social
compositions
(Tables
1 57
1 27
1 30
f
social
9.4 and 10.4).
distribution
chi
4.34
6.16
f1149 153.66
1 70 71.69
71.19 9.36
71.19 9.36
10.6
chi
0.14
0.04
1 36 47.43 2.75
1'76 103.13 7.14
1 25
43.37
7.78
0.62
0.44
3.09
1 16
1 16
I6
18.45
18.75
4.10
0.33
0.40
0.88
51.34
23.75
21.79
I
of adult
males.
groups
groups
and
foraging
Among the more important
following:
exp
79.50 10.22
were composed entirely
and foraging
of
84.11
140.40
Social
lobs
I
f
I
different
exp
in
I More Females
f
I
obs
The
multinomial
One Female
I
10.2.3:
found
of Three Age-Sex-Classes
Groups of 9 Compositions
No Females
No Males
198.7
177.5
62.9
1 178
1 70
441
frequencies
distribution
exp
I1 193
I1
1
es
Males
I obs
exp
obs
:ous Numbers
from multinomial
calculated
I
V
Fe
groups
were
results
were
10.4:
Table
Social
found in
of Three Age-Sex Classes
Compositions:
Various
Summary Table
Animals
of
Number of
the marginal
This Table represents
I
Age-Sex No Adult Females
I obs
Class
chi
exp
I
FI
---1
262
Y1
0.00
125.26
126
M1
I
I One Adult Female
i obs
exp
chi
I
I
I More Adult Females
I obs
exp
chi
I
1 154
I
122.53
8.08
No Adult
1 249
9.35
241.69
One Adult
Males
65.88
52
2.92
4.33
0.55 1 241 275.54
189 179.05
216.95
11.3
of Table
totals
Ci
1 101
0.22
2.77
119.16
More Adult
Male
Males
III
Mi---1
F;
Y1
214
334
237.77
328.18
178
1 173
210
2.38
0.10
177.49
174.32
221.14
1154
I 43
1 68
0.00
0.01
0.56
2.33
0.01
6.08
136.18
42.5
50.48
:II
1)
groups were less
social
while
groups
social
frequency
9.2)
foraging
(Table
3)
groups
than
frequently
while
a frequency
df = 1;
subgroups
a
with
ns)
(Table
less
observed
df = 1;
4.94;
squared =
Groups:
tendency
is
p<0.05)
P.
of
were
the
free
no adult
contained
female
no adult
9.2),
(chi
female
occurred
more foraging
squared = 4.94;
10.2).
Discussion
apparently
moved through
groups containing
equal
(Table
DeVore 1961, Altmann
individuals
male were
to chance (Table
nearly
p<0.05)
Social
The
social
than expected
groups
10.2.4:
no adult
(chi
expected
df = 1;
squared = 0.16;
containing
p<0.05).
10.2).
By contrast,
with
chance
to contain
male were observed
no adult
(chi
groups
df = 1;
squared = 4.83;
containing
to
close
than were foraging
male (chi
more than one adult
2)
likely
praplo
Mt. Assirik
at
shared
by
other
1979, Kummer 1968a).
socialising
to
interact
troop
without
10.7
to
Papio
socialise
troops
in
small
(Washburn
and
The manner in which animals
indicated
constraint,
a
society
but
in
in
which
which
there
between certain
was attraction
About 31% of all
but no adult
not
P. papio
that
exclusive
mating.
Although
the
were
associate
group.
male
and
one
by
10.3:
or
males
form
the
(Nagel
relationships
While
adult
together
that
groups
than
frequently
but
containing
10.3.1:
Nearest
At intervals
animals
restriction
vegetation
with
found
behaviour
is
than
more
to
females
adult
containing
1980),
one
frequently
as
as
presumably
not
from the analysis
1965b,
foci
the
P. hamadryas,
of
social
1972, ),
Rowell
Ransom and
In
periods.
social
form
infants
(Seyfarth
and P. ursinus
in
foraged
apparently
in social
1977),
for
especially
groups,
together
while
grooming
In
one adult
P. hamadryas,
into
one-male
groups
more than
one adult
male
Neighbours:
P. hamadryas in
occurred
male
teams
two
times,
apparently
more
forage
and social
rare.
Methods
of 20 minutes
adult
these
likely
male
in social
are
less
and were
they were unlike
more than
expected.
was likely
during
classes
young
containing
were present
that
tended
1971).
segregate
groups
age-sex
focii
males
to be found
social
no other
were
with
or more adult
females
adult
systems can be derived
(DeVore
(Altmann
P. cynocephalus
females
Such
on mating
P. anubis
in
one,
societies
Introduction
females
adult
behaviour
none,
were
groups
adult
between various
particular,
together
more
Neighbours:
of proximity
when there
only
in
were
and hence
group,
live
by chance,
expected
alone.
Some information
adult
containing
females
adult
a harem society.
of
Nearest
groups
of
that
not
female
one adult
a one-male
did
Furthermore,
chance
characteristic
within
Mt. Assirik
males
the
expected
only
those
in
already
adult
to
adult
with
at
numbers
close
indicates
observation
interact
to
suggests
females
This
at least
groups contained
social
male.
constrained
individuals.
animals
target
to hide
during
social
were selected
individuals
any of the three
10.8
periods
were
at
when more than 10
random,
chosen
animals
so
nearest
with
that
the
no
to them.
a
Figure-10.1:
Percentage
was nearest,
to an adult
in
age-sex
class
a given
which
observations
neighbour
nearest
and third
nearest,
second
during
female
periods
social
of
50U2
0
0
in
25
0
4-,
Adult
Males
Adult
Females
"-----------
Infants
Subadults
and
Juveniles
0
U
C)
P4
12
Rank
Dashed
in the
I
12
3
of
Neighbour
line
represents
population
3
1
(1 closest,
'proportion
.2
of
3
123.
3 furthest)*
that
age-sex
class
Figure.
10.2:
Percentage
was nearest,
to an adult
in which
observations
a given
age-sex
class
and third
second
nearest,.
nearest
neighbour,
during
social
periods.
male
of
50-1
Adult
U)
O
O
Females
O
En
A
0
Infants
w
0
Subadults
and
Juveniles
U
O
a
Adult
Males
123123123123
Rank of
Neighbour
Dashed line
represents
in tl, e population
(1 closest,
proportion
3 furthest)
of ' that
-age-sex
'class
The
age
and
sex
together
with
their
Another
target
the
of
three
nearest
distances
relative
from
was chosen
the
to
animals
from
this
target,
the
remaining
individual,
were
animals
recorded.
the
and
process
repeated.
The resulting
from
expected
data
the
were tallied
proportion
the
and compared with
the
of
various
age-sex
frequencies
in
classes
the
population.
The
frequencies
with
recorded
among any of
compared
with
10.3.2:
data
Nearest
For
Infants
(9.5%)
p<0.05)
but
closest
that
adult
females,
in
female
by chance
alone
(chi
(chi
= 5.99;
squared
p<0.05
Subadult
the
three
nearest
limits)(Fig
occurred
to
that
males
10.1),
expected
females
more frequently
adult
less
than
than
6.6;
=
= 1;
to
an
of
were
squared
= 3.98;
rather
neighbours
with
Fig
the
animal
df
1;
=
10.1.
to
expected
binomial
less.
in
an
because
closest
(95%
to
class
the
than
male
from
closest
that
in
of
classes
principally
summarised
are
= 1;
df
ns).
was the
be
to
and chi
adult
infants
= 1;
p<0.01)
not
males on fewer
age-sex
df
class
more likely
nearest
by chance
By contrast,
adult
three
were
signigicantly
occurrence
p<0.025
were rather
while
among the
df
These results
animals
not
the
tended
= 1;
respectively).
animals
= 4.28;
squared
deviate
age-sex
to
= 14.4;
df
nearest
among the
did
any given
males
(chi
squared
proportional
subadult
and
adult
and
(1973).
to adult
animal
distribution
the
squared
three
(13.9%)
males
(chi
population
calculated
were
and Dunbar
the
closest
expected
that
were
295 were made.
adult
was not
neighbours
of
to
The probability
adult
records
as the
general
animals
expected
nearest
classes
age-sex
Results
306
than
various
by Kummer (1968a)
given
were found
occasions
the
three
males,
for
made, while
the
Neighbours:
adult
the
which
be among
confidence
Adult
a probability
females
similar
alone.
males
were among the
expected,
while
(Fig
10.2).
expected
10.9
three
other
nearest
adult
neighbours
females
of
occurred
10.3:
Figure
frequency
which
with
from
deviation
expected
Relativ
female
to
adult
neighbour
are nearest
classes
age-sex
three
species
.
Exp)
(Obs-Exp)t
as:
deviation
(Relative
calculated
Juvenile
Adult
various
in
p_...
Female
.....
. Infant
,""o
t-Q
Adult
Male
1'ö
Subadult
-"8
.
-"6
Relative
-"4
Deviation
-"2
from
0
"2
Expected..
®.
"
P.
P.
T
Figure
10.4:
Relative
deviation
from expected
frequency
with
which
various'age-sex
classes
are nearest
to adult
neighbour
male in three
species.
(Relative
deviation
(Obs-Exp)
calculated
as:
+ Exp)
Subadult
Adult
o..........
J
--..
.......................................
Female
O""
----"""
-'89
Ö7 A-----------------
Juvenile
Adult
Male
Infant
-"4
-"2
Relative
0
Deviation
"2
from
"4
Expected
"6
papio
hämadryas
ZZ; elada
10.3.3:
Nearest
Adult
females
adult
females
those
for
animals,
neighbours
females
adult
in
frequently
that
the
three
nearest
neighbours.
groups
with
associated
of
female
upon
in
are
the
by
expected
female
company
are
the
nearest
three
the
three
nearest
expected.
Adult
have
to
three
and
grouping
to
similar
data
adult
males
This
alone.
bonded
a social
also
suggest
less
rather
it
makes
adult
nurseries
with
These
an
suggests
the
consistent
of
both
animals.
juveniles
This
groups.
chance
groups
social
kin
both
is
and
adult
closest
young
baboons
savanna
females
improbable
males,
as is
and P. hamadryas are rarely
found
socially
to
adult
case. in T. gelada.
The adult
in
their
have
for
species
expected
the
amongst
with
these
than
than
to
than
the
likely
more
tended
based
P. papio
amongst
P. papio
troops
organisation
found
female
amongst
reported
male
other
those
with
in
adult
frequently
much more
were
both
of
markedly
strongly
In
adult
female
social
were
neighbours
contrast
less
and
another
nearest
data
10.3).
females
males
and another
female
These
(Fig
have
T. gelada
T. gelada
of
Adult
that
to
adult
as the
1973)
1968a)
unlikely
of
male
females
(Dunbar
(Kummer
and
to be found
juveniles.
of
T. gelada
were
Discussion
tended
and
P. hamadryas
males
Neighbours:
the
males of both T. gelada
company of
associate
with
other
other
adult
adult
males,
whereas P. papio
as they
in proportion
males roughly
males apparently
in
occur
the population.
In
that
the
summary,
the baboons there
of one-male groups
10.4:
a troop
of
animals
species-specific
species
males
(Kummer
tended
to
Mt. Assirik
within
T. gelada
suggest
the constraints
or P. hamadryas.
Introduction
suggests
1980).
from
socially
to those of either
distance
(Schulman
tend
not interact
Distances:
Inter-individual
in
did
similar
Inter-Individual
data
nearest-neighbour
Furthermore
to
space
themselves
and
which
reflect
1974b).
Nagel
be found
much further
(1971)
10.10
between
relatedness
the
form
the
distances
distributions
social
was able
from
at
adult
individuals
which
are
of
the
organisation
to
show that
females
pairs
which
during
P. anubis
social
10.5:
Figure
Distribution
sex
200,
of
between-adult
distances
members
of
opposite
Observed
0
Negative
Binomial
'5
k=1.0
p=0.
t
C)
1o
C)
A
A
A
A
A
<1
Distance
123456789
Between
10
Neighbouring
Adults
of Opposite
Sex
(metres)
Figure
10.6:
Frequency
distribution
females during
social
100
of distances
periods
between
nearest
adult
n
v
a)
a)
50
<1
Distance
12
Between
345
Neighbouring
67
Adult
89
Females
10
(metres)
Figure
10.7-:
Distribution
of
distances
between
so
adult
;'':; males
Observed
0
Negative
O
Binomialk=1.5
p=0.5
0
C)
C)
vv
13
w
.
a
eee
<1
Distance'
12
3456789
Between
Neighbouring
10
Adult
males
(metres)
i
hkl
behaviour
than
three
all
he
troops
to
close
other
two
other
found
between
systems
of
P. anubis
towards
adult
females
in
10.11.1: Inter-Individual
least
When at
as
florescent
orange
the
where
areas
male
adult
or
frequently
secondly
troop
the
was in
the
the
not
were
were
baboons
both
rather
mating
restrictive
intolerant
of
males,
were
the
Identical
These
ignored
by
baboons.
the
10m from
introduced
seeing
animals
during
social
periods
this
arbitrary
If
target
because
ten
than
caused
cut-off
to
nearest
no record
vegetation
metres
apart,
distances
inter-individual
less
in
ground
the
animal
more than
with
helped
markers
firstly
(typically
Mt. Assirik
and
the
marked
on the
placed
male
were used with
methods
were
the
one of
adult
Bamboo poles,
me from
at
from
visible
nearest
observed.
was
adult
metre,
were
restriction
because
were
nearest
intervals,
was more than
small
the
individuals.
metre
were
to
which
female.
adult
and
chapter)
3 of
to
random,
at
prevented
this
Methods
tape
female
were generally
in
Distances:
it
reflected
males
males
be found
P. hamadryas
suggested,
P. hamadryas;
to
than
In
males.
unlikely
was much greater
but
target
the
This
was made.
and
the
P. anubis,
the
distance
estimate
at
visible
nearest
females
adult
were
in
he
distance,
the
chosen
males,
and the
males
10 animals,
I estimated
adult
that
findings,
and
males
x P. anubis
in P. hamadryas.
each other
visible,
adult
but
adult
These
troops.
P. hamadryas
or
that
males,
adult
distance
typical
P. hamadryas
were
data
one metre;
few
data
be
to
rejected.
10.4.2:
Inter-Individual
In 95% of
within
5m of
the
Distances:
observed
the nearest
cases they were less
cases (n: 386),
adult
animal
than 1m apart.
less
than
a metre
less
than
5m away on 91%.
metre of another
Results
away on 29% of
adult
female
Similarly,
was sitting
In 58% of
sex.
males had another
observed
adult
in 55% of cases,
10.11
animal
the opposite
of
Adult
all
an adult
occasions
females
and within
adult
male
(n=205),
and
were within
5m in 96%.
one
Figure
10.8:
Distribution
opposite
sex
between
periods
of distances
during
social
100
a hybrid
0 anubis
"
adults
nearest
of
papio
hamadryas
ETp
0
cl
ö501
4-4
1ý1i ýI 1¢ýeä
T
a
<1 <5 <10 >10
Distance
Figure
Between
<1 <5 <10>10
Neighbours
of
- <1
Opposite
<5 <10)10
Sex
(metres)
10.9:
Distribution
of
distances
between
adult
in
males
three
100
0
41
ce
r7
C)
r0
°
50.
0
c
v
v
t,
aý
a
<]' <5<10 alp
Distance
Between
<1 <5 <10 X10
Nearest
Neighbouring
-1
<5 <10 ;, 10
Males
(metres)
species
The
distribution
the
members of
chi
8.52;
=
between
distances
data,
df
is
in
the
10.9).
P. papio
P. anubis
might
troop
and
differs
statistic
In
adult
the
in
females
P. papio
to
be
found
those
troops,
and especially
10.4.3:
Inter-Individual
baboons
These
social
or
leading
data
to
with
differences
partly
in
activity
social
between
The data
this
it
may
have
(Schulman
1980),
adult
male P. papio
male.
These data
is
)
males
in
P. papio
the
of
any
and
and
in
numbers
but
other
baboons
baboons
of
in
the
not
be
poor
troops,
distances
(1971)
Nagel
to
response
a
of
their
of
Inter-individual
since
should
did
a reflection
been
areas.
this
here
also
for
(1971)
were likely
are
value.
test
also
responsible
took
his
for
the
two studies.
by Nagel
presented
large
social
presented
those
the
the
the
the
adult
than
together
closer
that
periods
in
null
when
males
adult
males
probable
due to
the
Discussion
sit
although
crowding
during
to
seems
(Fig
P. harnadryas.
of
tended
the
did
than
Distances:
organisation,
visibility
vary
It
species.
other
together
closer
but
df = 2;
P. hamadryas
in
found
those
hybrids,
between
in
troop
respectively)
critical
distances
of
by Nagel
accepting
nominal
adult
P. anubis
indistinguishable
the
resembles
x P. anubis
P. hamadryas
tend
distribution
the
summary,
from
2;
=
between
6.50;
=
squared
against
are
slightly
x
p<0.09
warns
distributions
only
= 2;
df
p152,
chi
df
found
those
P. hamadryas
test;
Assirik
= 45.2;
distances
from
of
(1971)
Mt.
at
squared
of
differed
6.24;
=
found
chi
hybrid
the
1977,
two
that
hypothesis
in
squared
(Pollard
10.5).
test;
two-sample
chi
that
test;
Nagels
to
according
from
have
also
and
(Kolmogorov-Smirnov
p<0.08
The distribution
10. $).
distributions
The
from
(Fig
P. hamadryas,
adult
different
significantly
two-sample
two-sample
(Fig
is
between
(Kolmogorov-Smirnov
different
significantly
p<0.001)
P. anubis
p<0.05)
in
males
(Kolmogorov-Smirnov
males
= 2;
(1971)
by Nagel
given
Assirik
Mt.
at
squared
sex in
opposite
found
that
distances
of
P. papio
for
are
clearly
P. hamadryas,
to be found within
different
10.12
from those
different
especially
5m of another
presented
from
because
adult
by Nagel
on
P. anubis,
since
found close
hybrid
to an adult
The observation
this
more likely
rather
they
respect
rather
P. hamadryas males tended
that
their
reflects
into
segregation
males in P. papio
between adult
association
in
male;
were
be
to
the
resembled
Nagel studied.
baboons that
presumably
P. papio
female
adult
to avoid
one another
one-male groups.
The close
makes their
and may indicate
one-male groups improbable,
relationships
genetic
close
into
segregation
between them (Schulman 1980).
10.5:
Male
and Female
(1971)
Nagel
the
on
1)
for
whether
adult
2)
whether
adult
female
baboons
they
belonged
the
is,
that
provides
with
in a detached
with
to
back
followed
check
whom
party.
female
is
after
he
Adult
females
in
P. anubis
females
do
the
that
the
male shortly
adult
he had gone 10m.
before
nearly
P. hamadryas,
males do not.
female
moved away, specifically,
P. hamadryas troops
or sitting
look
to
differing
Nagel
troops.
to
back when moving away from an
male glanced
tend
to
led
including:
behaviours,
whereas P. anubis
or not
structures
x P. anubis)
been interacting,
P. hamadryas males
following,
mating
whom he had been "associating",
with
he had either
and
on whether
or not the adult
female
male
adult
'indicator'
three
in
differences
(P. hamadryas
hybrid
or
Introduction
that
depending
counts
P. anubis,
data
showed
between
interactions
several
Interactions:
follow,
always
while
not.
3)
whether
away from
an adult
P. hamadryas
back,
10.5.1:
the adult
or not
while
female
the
in P. anubis
female
observe
the
position
of their
and the
female
walks
rarely
initial
paid
of
both
In
without
checking
seen.
Methods
these data were recorded
movements
off
is not usually
such behaviour
Male and Female Interactions:
At Mt. Assirik
to move
whom she had been associating.
male with
groups,
back when starting
glanced
individuals
If
association.
no attentiong
10.13
or
if
whenever it
over
was possible
to
10m from
the
about
the male moved out
either
began to
of
interact
sight
with
or if
anot.. er animal,
that
social
both walked off
likely
to
(19%).
occasions
occasions
When females
(Table
(13%)
look
back.
left
first,
to
and were thus
in
:tion
In
p<0.05).
observed
86 occasions.
of
interac
an
= 1;
df
never
were
males
terminate
= 5.63;
squared
I assumed
Results
on 54 out
first
Females moved off
more
directions,
had been broken between the two.
contact
Male and Female Interactions:
10.5.2:
(chi
in different
this
sample
32 leavings,
their
Females
they
adult
followed
on
6
on
6
back
checked
10.5).
Table 10.5:
Behaviour
Adults
of
on Ending
Behaviour
YES
Male checks
back
Female checks back
Female
follows
Obedience
In
out
responses
the
closely
(1971)
Nagel's
of
86
1: 8 ratio
terminology,
Mt. Assirik,
lacked
this
respect
Awash.
herding
behaviour,
groups
maintained
apparently
behaviour
suggest
and that
by the
32
6
48
54
6
26
32
12
75
86
In
they
this
10.14
resembled
and were divergent
and P. hamadryas.
and adult
indices,
typical
at
females
of
in
showed that
P. pap
closely
Mt. Assirik
with
the
adult
and that
in
P. anubis
at
P. papio
males;
showed no
to live
in
at
baboons
these
of P. hamadryas.
these baboons were unlikely
behaviour
"disobedience"
Discussion
corresponded
that
75
gave
x P. anubis
males
as measured by Nagel's
These results
32
of Ppapio
between adult
their
0
by Nagel in P. anubis
observed
the coordination
n
(1: 6).
Male and Female Interactions:
Interactions
NO
females
opportunities
from the 2: 1 and 11: 1 ratios
10.5.3:
an Association
in harem
short,
they
to possess a mating
were unlikely
10.6:
all
social
is
thought
primates,
in
particular,
(Sade
likely
to
in
males
females
their
frequency
individuals
in
infants,
who
observed
in
frequently
females
of
P. anubis
troops
and
form
centres
in
groom
adult
other
one-male
The distribution
of
the
leaders
do not
approach
least
1.5m
(1968a
He goes
(the
"leader"
1968a,
p55).
P. ursinus
If
each
other
keep
"adult
on to
males
and "follower")
has not
P. papio
in
between
adult
adult
males
free
their
adult
male
this
with
has
hand,
females
distance,
but
often
"the
unit
ordinarily
Kummer
his
groom
"Thus
other:
leader"
each
groups
one-male
groom each
P. anubis,
also
mingling",
within
with
1963).
is
males
while
males
the
with
for
from
been
DeVore
and
interact
no females
only
other
females
P. hamadryas
rarely,
Females
Kummer and Kurt
units
having
these
of
with
to
to
1968a,
grooming
at
from
groups.
other
species,
grooming
(Kummer
P. cynocephalus
and
been reported.
lives
in
one-male
we predict:
a)
this
expected
Washburn
the
on
not
may also,
engaged
Grooming
that
mention
(see
He
groom
second
activity;
adult
of
to
apart...
follower...
adult
between
structure
that
notes
p47)
other".
the
social
are
than
adult
to
interacting
baboons
(Kummer
units
grooming
to
at
but
troops.
attracted
P. hamadryas,
males,
related
keep
In
from
social
savanna
of
prep).
of
hybrid
three
the
within
groomed
the
the
are
therefore
were seen
often
social
males
females
of
males
troops
the
He related
adult
many
in
or
more
organisation
by
adult
adult
population.
social
then
Collins
P. anubis
is
system
4).
many
of
relationships
that
significantly
in
and maintenance
grooming
group
(Chapter
time
consequences
mating
showed
P. anubis
the
the
of
in
not
social
The
(1971)
infants
restrained
not
1961,
the
in
to
patterns
but
in
with
observations
of P. hamadryas.
up most
establishment
1977).
Nagel
P. hamadryas
adult
are
the
important
the
Seyfarth
example,
showed that
also
in
influence
For
troop.
have
to
1965,
took
grooming
activity,
Grooming
bonds
that
Introduction
Grooming:
Of
system like
groom one another
10.15
groups
similar
to
those
of
T. gelada
b)
e)
adult
females
basis
of
adult
females
their
by other
If
more frequently
groom one another
in
abundance
the
infants
with
population,
are
than
and
more frequently
groomed
on the
expected
than
expected
females.
adult
P. papio
lives
in
one-male
groups
similar
to
less
expected
those
of
P. hamadryas we predict:
a)
that
adult
b)
that
adult
their
c)
that
males
females
the
females
adult
P. papio
groom one another
groom one another
in
frequency
more frequently
If
seen to
are
population,
the
resembles
do not
b)
adult
females
groom one another
c)
adult
females
with
age
infants
and the
results
of
the
sex
and
in
more frequently
than
expected,
groomed more frequently
are
the
active
pairs
were
of
grooming
all
(Chapter
tallied
than
(groomer)
partner
during
recorded
4).
When the
not
be determined
null
hypotheses:
could
partners
and
expected
and
each
age and
the
scan
sex
of
sample
one
was excluded
pair
the
of
or
from
analysis.
I tested
2)
then
females.
adult
recipient
1)
females
Grooming: Methods
The
the
groomed by adult
not
groom one another
males
other
from
particular,
baboons,
savanna
adult
10.6.1:
are
than
than expected.
a)
by other
in
and,
infants
with
rather
The
the
age-sex
following
classes
were
their
proportion
with
Given
each age-sex
that
member of
a grooming
recorded
as members of
abundance
in
class
occurred
the
pairs
in
population.
with
each class
pair,
grooming
frequency
a certain
was equally
likely
to
as a
groom or
be groomed.
3)
Given
the
tendency
each age-sex
Several
expected
the
class
studies,
frequency
proportion
of
chose
its
notably
with
of
each
those
which
class
to
partners
at
random.
that
Nagel
age-sex
by
age-sex
classes
10.16
in
classes
the
groom
or
(1971),
groom or
population.
be groomed,
calculate
are
groomed
This
could
the
from
be
as the null
stated
4)
that
the
hypothesis:
from
animals
that
groom any animal
The first
each
The
that
the
the
of
class
with
the
(of
(chi
the
of
(chi
the
which
for
animal
groomers
to
it
squared
age-sex
in
each animal
which
was grooming
goodness-of-fit)
classes
in
occurred
class
several
85% of
about
females
of
classed
(0)
juveniles
Independent
M.
pooled
(M).
The
classes
data
example,
samples)
Nagel
adult
males
the
multiplying
by
pooled
into
in
in
the
matrix
(1971).
were
The
seen
to
of
adult
corresponding
episodes.
For
this
ensure
that
matrix
had expected
values
of
with
adult
M.
order
pooled
were
Females
were
(S
Subadults
(Y)
Tr,
were
B1)
or
and young
infants
with
but
grooming,
tested
were
pooled
Z)
or
5
with
older
W.
all
pooled
were
classified
as
((M))
were
adult
males
follows.
as
ux5
the
then
groom
for
frequency
expected
adult
males
in
The
matrix
MFYxMFY
proportion
10.17
age-sex
matrix.
was
as before,
3x3
proportion
the
aüx5
hypothesis
null
first
further
(M)
given
to
juveniles
males
in
a given
in
groomers.
adult
in
pooled
of
(Br.
The expected
animals
animals
females
status
ages
resulted
fourth
(E)
all
sum of
matrix
two distributions
totals.
grooming
resulting
active
of
Mature
were
the
younger
and
This
were
as recipients
as
infants
infants
classes
and oestrus
(Fi)
(F)
females
in
R_
sum of
total
reproductive
unknown
separately
other
C=
T=
and
(P)
resulting
up an nxn
the
with
marginal
where
grooming,
cells
pairs
the
giving
age-sex
the
participants
the
comparing
by setting
grooming
as RxC/T
Pregnant
more.
of
grooming
actively
analysis
was tested
and recipients)
receiving
and
the
goodness-of-fit).
frequencies
class
by dividing
recipients
hypothesis
were calculated
age-sex
was tested
and
squared
null
observed
values
hypothesis
null
The third
the
likely
equally
by tallying
was compared
groomers
active
distributions
still
were
population.
into
or
age-sex
frequencies
The second
of
was tested
distribution
resulting
with
the
class
class
they encountered.
hypothesis
null
age-sex
each age-sex
females
age-sex
was collapsed
comparison
with
which,
was computed
the
population
of
adult
(of
females,
with
for
by
scan
and
this
multiplying
10.6.2:
age,
the scan samples 650 grooming
and where
grooming
various
10.6:
Observed
Age-Sex= PropClass = ortion
I
E
Fi
S
Z
B1
(J
recipient
distribute
(chi
their
proportion
and juveniles
sex classes
classes
age-sex
50.46
chi
squared
10
4.8
025
.
031
.
52
21
32.6
39.7
-
11.50
8.77
4
2.6
-
158
97
24
183.0
165.0
66.2
3.33
28.09
26.88
3
6.2
1.68
19
21.2
0.23
20
37.1
7.85)
9
11.1
0.38)
1300
1299.25
002
1 005
.
1 061
.
009
classes
did
not
grooming
equally
participate
p<0.001)
among the
randomly
df = 12;
received
140.09
as
(Table
groomer
and
10.7).
to groom and be groomed the baboons did not
tendencies
the available
(Table
various
oomin
382.0
p<0.001)
males and juveniles
to
10.6).
p<0.001)
004
squared = 47.7;
Adult
(Table
522
squared = 102; df = 13;
Given these
squared
0.62
.
in
population
0.30
1
(chi
of
be
(chi
17.7
.
The age-sex
the
df = 13;
could
participate
330.0
1 029
.
1
(I
in
the
not
340
1 141
.
1 127
.
1 051
.
Tr
did
21
.
0
Y
Br
seen,
in which
participants
Expected
254
1
P
both
of
Observed
grooming
.
I 295
.
014
.
F
occurred
frequencies
and expected
M
episodes
were recorded
classes
squared = 140;
chi
pairs
sex
age-sex
as they
in proportion
goodness-of-fit;
the
appropriate,
The
determined.
Table
number of grooming
Results
Grooming:
In
by the total
product
did
grooming
(Table
not distribute
grooming
partners,
disproportionately
10.8).
10.18
available
age-sex
classes
grooming
time in
10.8).
their
while
both
from
the
adult
females
various
age-
Table
10.7:
Frequency
of Grooming and Receiving
of Observations
Various
Age-Sex Classes
Age-Sex
Groomer
M
117
F
chi
223
squared
33.05
318
204
24.90
E{
6
15
3.86
P
2
8
3.60
23
29
S
Z
4
2
0.69
17
2
8.05
0.00
0
95
63
6.48
Y1
53
44
0.84
10
14
0.67
0
1
3
18
15.21
Fi
Br
1
;
Tr
B1
Table
Recipient
Grooming
(J
.
15
5
5.00)
(I
i
4
5
0.11)
650
650
102.46
10.8:
Contingency
Table for Grooming Partners.
Expected values are given from marginal
totals
GROOM
RECIPIENTSI
M
ERS
F
J
I
MI
obs 1 42
1 exp I 40.1
133
119.7
40
58.0
8
5.1
FI
obs 1 56
1 exp 1 40.9
101
121.9
67
59.0
3
5.2
Fi
I obs 1
1 exp 1
JI
obs I
4
21
4
0
5.2
15.6
7.5
0.7
11
exp 1 23.6
II
obs {
I exp 1
4
7.2
10.19
66
52
2
70.3
34.1
3.0
28
21.5
6
10.4
2
0.9
b
interest
Of particular
in
males
(Table
Adult
10.8).
(chi
expected
females
significantly
Grooming
p<0.001)
(chi
43.7;
=
squared
df
partners
more
10.8).
Adult
but
expected
than
not
ns).
distributed
among
evenly
Adult
females
groom
far
more
squared
= 258;
females
adult
frequently
and chi
the
squared
age-sex
df
both
= 1;
with
than
expected
= 55.8;
df
= 1;
10.9).
10.9:
Table
Contingency
Expected
values
are
table
for
given
from
grooming
M
RECIPIENTS
M
F
; obs =
42
1 exp ;
41.9
1 obs 1
56
1
exp
i obs i
1 exp 1
J
;4
obs
exp ;
I;
10.6.3:
Grooming:
Grooming
reminiscent
of
I
40
8
13.4
67
101
3
66.7
68.8
4
0
5.3
1.3
21
5.5
16.5
11
66
52
54.2
72.1
69.9
17.3
13.4
28
17.8
62
17.3
4.3
2
Discussion
relations
showed
J
54.2
55.8
4.1
population
RS
133
4
exp 1
in
F
51.7
1
9 obs
Fi
partners.
proportion
G ROOME
males
than
df = 1;
p<0.001
females
(Table
more
groomed adult
grooming
adult
(chi
= 1;
males
p<0.05)
infants
the
respectively)(Table
p<0.001
groomed
population
in
10.9).
adult
as available
df = 1;
not
infants
without
and
also
with
are
occur
(Table
study,
squared = 1.89;
partners
they
as
males
females
(chi
so
this
occurred
squared = 5.61;
groomed
classes
they
as
proportion
to
in
P. papio
considerable
that
exhibited
at
affiliation
by
10.2 0
Mt. Assirik
to
suggested
each
P. hamadryas
or
that
other,
T. gelada
adult
strongly
males
I
the
from
excluded
the
one another
were
the
adult
same time,
females
adult
other
social
organisation
males,
to
consistent
in
with
the
male,
breeding
with
in
adult
in
in
as
groups
which
in
which
not
are
They
free
adult
are
with
Nor
they
are
seen
consistent
by an adult
from
are
excluded
also
consistent
with
a
adult
normally
are
each
the
with
other
a
and
for
affiliation
show strong
males
with
by
"adopted"
interact
to
At
P. a nubis,
females,
They
males
who
interaction
behaviour
are
male
males
consistent
adult
females
adult
population.
P. hamadryas.
some adult
are
the
occasions
T. g elada,
societies.
T. gelada.
of
of
in
affiliative
of
an
breeding
of
in
baboon
females
and
males,
males.
adult
Summary
Exclusivity
is
observations
of
female
time
every
mating
of
group
social
not
adult
she
(Strum
pers
In
savanna
comes
into
oestrus
with
her
and
T. gelada
the
male
has tenure
entire
defend
his
females
In
receptive.
known adult
However,
with
males
from
to
order
on one-male
society
baboons
males
it
groups
is
unlikely
a mating
would
have
his
system
like
been
found
P. papio
that
in
10.21
of
all,
pers
the
time
for
the
male
must
of
it
the
and
are
has
system
all,
to
cycles.
Mt. Assirik
nearly
sexually
a mating
many oestrus
at
not
would be necessary
P. hamadryas.
or
comm, Lee
rest
they
P. papio
through
many
favourite
persists
group,
when
even
its
references).
Boese 1973,1975)
that
the
much of
over
males and females
pens
in
are
a
consort
exclusivity
that
establish
who
other
this
males
other
(cf
There
(Collins
for
females
over
species-specific.
P. hamadryas
that
follow
entirely
comm, and see page 10.2 for
time
based
by one male baboon
rights
associate
comm) and may also
adult
groups,
as in
adult
which
data
coercion
of
the
found
These
and
some
a freedom
that
upon
kin
in
from
to
savanna
which
least
at
showed
low frequency
perhaps,
other
particular
10.7:
harem
population,
society
females
equivalent
males
adult
and,
that
two
on one or
female
adult
excluded
based
live
a society
with
not
an
and P. ursinus.
P. cynocephalus,
between
grooming
suggests
which
consecutively,
However,
population.
males
adult
observed
groomed
mating
lived
in
In P. hamadryas
social
groups.
a
About
a third
of
adult
females
but
males
such
those
unlike
to
tended
sit
the
close
and the
P. hamadryas
genetic
T. geladaa
nearest
as
P. hamadryas
to
other
savanna
1980).
responsible
for
"disobedient"
Adult
This
the
females
maintaining
as P. anubis
observation
suggests
in this
that
that
in
did
females
affiliative
males
10.22
but
to
similar
differ
from
the
individuals
in
as
adult
frequently
sexes
T. gelada
a society
behave
was
like
both
or
the
both
with
groups
though
they
males,
being
were
as
males.
to have emerged from this
behaviour
society.
animals
may indicate
towards
results
adult
adults
apparently
to
were
adult
of
age-sex
This
not
proximity
entirely
societies,
huddles
they
between
One of the most interesting
is
P. paapio
baboons.
relationships
(Schulman
this
of
contained
The
seen.
T. gelada
or
study
composed
never
neighbours
adults,
In
this
groups
were
baboons.
savanna
when socialising.
clusters
close
in
found
in
found
those
in
found
in
observed
Social
males.
found
animals
groups
social
no adult
as those
of
classes
the
groomed one
study
another.
among the males was important
Organisation
CHAPTER11: Ecology, Social
in the social
Much of the variation
to
The diet
ecology.
organisation
social
the
ascribe
cannot
possible
seasonally
large
wet
that
there
baboons.
few
animals
source
at
groups
within
animals
has also
Diet
finding
food
enough
seasonal
fragmentation
the
of
frequency
the
movement and
by
baboons.
the
much of
the
suggests
that
largely
That
behaviour
the
of
baboons
visited
enter
such
of
of
troops
study
Since
the
field
This
of
Papio
occurred
ni
the
same food
of
small
foraging
encouraged
the
the
ranges
to
some of
a crude
method
baboons
these
the
cover,
habitats
they
were
11.1
further
in
parts
of
the
was capable
in
is
distributed
even when
both
of
speed
were used
predicting
so
remarkable,
and
ranges was determined
visibility,
moved across
in
variation
range
of
itself
in
evidence
habitat,
changes
area was made up of
and which
baboons.
non-existent
seasonal
they used their
in canopy
or
gives
show that
various
which
these
of
small
for
compensated
for
ranging
have had some difficulty
and
day
was able
with
differences
About a third
them.
length
the
would
ranges,
the manner in which
by local
rarely
troop
the
accounted
classified,
coarsely
the
The present
productivity.
influenced
smaller
in
difference
so
from
may have
turn
season,
the troops.
that
in
in
in
the
group.
normally
formation
the
which
apparently
suggest
to
dry
the
feed
to
ambush
change
population
animals
able
led
troop,
within
Long day ranges
these
of
was
meant that
social
other
one
by
seasonal
the
of
it
food
predation
in
the
T. gelada,
because
apparently
together
only
normally
This
moving
to form cliques
in
food
were
same time.
the
forage
known
the
since
the
of
strong
fragmentation
is
Thirdly,
trees,
risk
Secondly,
not
seasonal
a
was
fragmentation
seasonal
that
could
groups
season
when
form
to
able
we
in
groups
as we can for
the wet and the dry seasons
between
productivity
time
a
highest.
probably
was
predators
fruit
at
superabundant
were
although
large
in
on its
effect
Firstly,
survive
of grass,
troops
on demography,
effects
reasons.
to
solely
large
these
that
three
of P. papio
consisting
of baboons has been
has a profound
probably
least
at
ability
to a diet
wet season
is
for
its
through
P. papio
of
System of P. papio
organisation
or indirectly
directly,
either
related
and Mating
and productivity.
of habitats
rapidly
which
the
when they
did
as a mosaic
throughout
led
lacunae
to apparent
within
was large,
area ranged over by one troop
about a third
was probably
The position
of
the
In
used
by the
baboons
tree
in
field
the
reaches
forking
trunk,
their
often
here.
the
troop,
as
that
suggested
of
feeding
behaviour,
to
although
interaction
took
members of
troops,
the
there
troop.
place
of
In
such
probably
a society
it
11.2
is
force
troops
High
not
foraging
during
This
and
rather
in which
individuals
are
for
conceivable
in
permitted
in
social
peak times
that
most
social
between
than
of
that
may suggest
all
there
were as many
in
baboon
most
much substructuring
that
to
productivity
spent
observed
baboons
the
apparently
groups,
be opportunity
be
could
of large
form,
activity.
there
as
and
water,
of
to
subgroups
large
such
sex
to
necessary,
small
within
either
would
not
was
With
the troops.
animals
adult
contact
the
patterns
ranging
by baboons
seasons.
troops
in solitary
were engaged
social
extended
both
time
was apparent
sources
park
the
of
subdivision
on the
been
the individuals
of
most
Social
home ranges.
may have
of
much
spend
behaviour
social
allowed
which
season,
the
parties
sleeping
and offspring,
of neighbouring
large
topmost
from
roosting
formation
the
promote
in
the
in an area which was seasonally
sites
and moving
in
water.
of permanent
troops
of
thus
the
of
species
the ends of the branches,
permanent
areas
was
near
influence
of
various
large
of
spend much time
the
mothers
helped
have
may
One effect
baboons
of
which
sites
tree.
of
gallery
horizontally
isolated
the
number of sleeping
productive
wet
rather
location
the
and to encourage wide overlap
the
small
had a major
sites
usage
The limited
highly
of
by the distribution
predicted
until
this
only
shape provided
near
composed
did
area
by the geology
of
almost
grew
This
points
units
These sleeping
of that
pentandra,
branches
with
branches
ends.
Ceiba
Individuals
tall,
the branches
into
surface
supported
sources
tree,
of
night.
often
formation
throughout
scattered
at
These
near
the
encouraging
troop,
were
and at few other
the trunk
of
as refuge
tree.
the
of
the
part
determined
water
permanent
one species
area
the productive
was largely
water
the
turn,
and, in particular,
forest,
Although
smaller.
of permanent
area.
home ranges.
their
areas has
of such little-used
the home ranges of the baboons the existence
males
within
might
not
transfer
always
This
lead
might
found
those
out
to
further
socially
those
with
baboons
male
both
seen
in
as
within
the
troop,
of such intriguing
of
associated
the
it
to breed.
to
analagous
out of their
natal
must await
possibilities
Adult
the
differences
between
adult
males.
tolerant
of
Grooming
between
each
genetic
ties
others
three
among adult
mating,
A possible
structure
four
of
males
here
for
is
closely
kin
groups
other
Ecological
units,
to
these
such
or
social
foraging
units,
animals
one
another.
baboons.
further
to
associated
those
were
coalitions.
may have
had
Agonistic
of
soliciting
threatening
and perhaps
baboon
societies
of matrilineal
one or more adult
males
units
with
with
consistent
kin
a network
parties,
might
have
although
adult
males
11.3
among
study.
social
which
some
males
Such social
in
seen
however,
mutual
at Mt. Assirik
the troops
form
much
led
be unique
would
to
these
involved
sometimes
male
one in
were more or less
and with
and
and is worth
competitive
presented
with
often
males
adult
in
seen
particularly
that
of
behaviour
and probably
that
of
either
males
were,
baboons,
affiliations
males
in
by adult
There
evidence
all-
that
the
adult
Mt. Assirik.
relationships
adult
by
of adult
suggested
like
in
sex had a nearest
females
savanna
large
in
society
Finally,
females
consistent
be found
same as that
groups.
and the
males
threatening
of
the
and formed
between
groups
was not
other
interactions
between
either
was considerable
adult
found
not
of
adult
at
to
an animal
There
strong
support
observed
unlikely
expected
grooming
P. papio
particularly
in
of
intense
was not
males
one-male
coercion
were
were
The
mates.
parties
a
class
with
or the
T. gelada,
that
is
and
large,
with
for
competed
in
been
age-sex
species
with
baboons,
savanna
might
adults
Mt. Assirik
at
baboons,
savanna
and foraging
groups
have
the
the
which
of P. papio
system
in
seen
societies.
a given
P. hamadryas,
data
that
The probability
baboon
closely
transferring
social
harem
with
neighbour
groups
males
in
troops
of
T. gelada.
the
but with
and mating
to
similar
groups,
with
within
organisation
complex
composition
the
clans
remain
study.
was apparently
in
of
formation
the
The resolution
The social
the
but
natal
in P. hamadryas,
to breed.
clan
group,
their
of
throughout
the
corresponded
may have
troop.
fairly
foraged
on
L
11.1:
Figure
possible
social
(This
figure
is
of
structure
not referred
the troops
to in the
of P.
text)
papio
at
Mt.
smallest social unit families of females and associated,
familiar, mates
O
extended units
foraging
units
VNI
oý
"small"
dry-season
troops
(clans?)
large
wet-season
troops
Assirik.
their
own, with
the
social
unit
social
with
large
these
season
unit
of
P. hamadryas.
may have occurred.
differences
Genetically
P. cynocephalus
individuals
in
(Pollock
Mali
suggests
possibly
under
Baboons
the single
of
of baboons
find
food
increasing
into
lush
alpine
Simultaneously,
forest,
near
where
the
it
forest
and drills
these
not
floor.
animals
lack
into
terrestrial
These
a tail,
exploiting
found
the
which
tends
the
11.4
and
niche
to
papio.
to
desert
The
and
them
move
to
would
potental
niche
may have
given
radiation
competence
the
on
canopy,
the
but
all
the
belief
other
in
other.
deep
on or
Both
mandrills.
support
to
them
reinvaded
in the
which
in
or woodland,
adaptive
ancestral
from
baboons,
decreasing
not
were
savanna
allowing
probably
animal
foods
ground
led
savanna
P. papio
individuals
adaptations
one hand, where
and
began to exploit
mandrills
are
newly
the
have
may
this
meadows on the
the
with
of
enabled
Thus the wider
on
competence
P. anubis
as possible.
fringe
which
is
than
show facility
diet
their
such trees.
Finally,
disadvantage,
was no
in
1969).
and
subspecies
More omnivorous
trees.
individuals
to
the
with
et
containing
ecology
Mandrillus
an adaptation
between
new foods.
be included
and possibly
increasing
advantage
the
trees
scattered
fruit
by
made available
exploit
been
on the ground
P. papio
forest
have
been
may
may have
Troops
The behavioural
some
(Wiener
P. anubis
1973).
between
as much fruit
in
are
are in any case no
from
P. cynocephalus,
genus Papio
the
habitat
between
and Brett
species
include
terrestriality
(Jolly
and
between
units
and Moor-Jankowski
no more distinct
should
rather
met
and P. anubis
is
comm).
they
that
and seem to
original
easily
pers
closely
fragments
P. ursinus
and
dry
of any one
troop
social
with
the
other
season
wet
and
(Weiner
intermediate
appearance
further
trees
P. ursinus
or
of
groups
baboon
Guinea
the
with
in baboon species
among human races
those
than
blood
in
In
When these
of P. cynocephalus
those
associated.
most or all
the
of
P. papio
of
more than
with
individuals
groups
from
different
greater
exist
blood
the
a subset
both
of
animals
same fragment,
the
clans
fragments
1970),
probably
form
some movement
al.
in
generally
to
socialised
respects
were
units,
the
Although
they
together
immature
or with
fragmented,
groups
social
to
males
which
staying
associated
analogous
adult
other
that
baboons
but
radiated,
(Jolly
If
correct,
to
that
a secondary
in
opposite
Thus Theropithecus
adaptation.
from
directions
a Papio-like
1970).
this
then
in
correspondence
Assirik
is
theirs
differentiated
and Mandrillus
form
that
interpretation
in
the
the
ancestral
plant
genera
a habitat
baboons
and
habitat
ancestral
may be exploiting
P. papio
which
of
at Mt. Assirik
evolved,
patterns
of
of much of Africa
and in the Plio-Pleistocene
11.5
especially
habitat
is
baboons
of
similar
the
given
found
(McGrew et al.
at
Mt.
1981).
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