Psychological Warfare and the Management

of Relationships between Male Baboons

(Papio cynocephalus)

by

Carlos Drews

Dissertation submitted to the University of Cambridge

for the degree of Doctor of Philosophy

Department of Zoology,
University of Cambridge,
Downing Street, Kings College
Cambridge CB2 3EJ Cambridge CB2 1ST
._---_ .. __._.. _._--~....""
~BSTRACT OF DISSERT~TION,

T~BLE OF CONTENTS

~ND

~BSTRACTS OF CH~PTERS
 PSYCHOLOGICp~ Wp~FARE ~~ THE ~~~AGK~ENT OF RELATIONSHIPS
BETWEEN ~~~E BABOONS (PAPIO CYNOCEPaA~US)

~~STRACT OF DISSERTATION

Social animals derive benefits from living in a group, but at the same
time, they compete with each other over resources. The opportunity to
interact repeatedly with other members of the group is the basis of
their social relationships. Natural selection should favour individuals
that manage their relationships to their advantage. This study
addresses the role of social interactions between male baboons in the
management of their competitive relationships. Free-ranging baboons
were observed in Tanzania for a total of 1413 hours, distributed over
16 months. One advantage of studying male baboons is that the
development of their relationships can be monitored from the beginning,
when alien males immigrate into the study troop. The concept and
definitions of dominance were reviewed, because dominance is central
to the description of competitive relationships. A distinction was made
between the short-term, immediate circumstances that motivate a social
interaction and the possible long-term social goals associated with it.
Males were selective with respect to the target and frequency of their
interactions, in ways which suggest that they actively manage their
relationships through social behaviour. Gradual changes over time in
the responses to approaches within male dyads illustrated the dynamic
nature of their dominance relationships. Participation in coalition
attacks did not seem to influence dyadic male relationships. An index
of spatial intolerance was developed, in order to test the effect of
varying interaction rates on the pattern of responses to approaches.
The higher the rate of interactions received by a sUbordinate, the more
promptly it retreated upon being approached by the dominant. This
effect suggests that intimidation can be an important competitive tool
for male baboons. Management of relationships took place in the course
of mundane social interactions, \olhich were generally not overtly
aggressi ve. It was postulated that interactions between males have
elements of honesty, bluffing and assessment of the competitive ability
of the opponent. Relationships between male baboons are best described
as a case of "psychological warfare". The basis for psychological
warfare is the high risk of injury for both males, should the
interaction develop into a fight. In male baboons, injury inflicted by
their canine teeth is known to result in reduced competitive ability
and sometimes even death. It is suggested that the main function of
relationship management in males is to reduce the risk of escalation
in subsequent encounters. The results affect our perception of social
interactions, dominance and competition between animals in general.

1
 TABLE OF CONTENTS

PAGE

ABSTP~CT OF DISSERTATION
1

CfI..J>..PTER ONE

"RELATIONSHIPS, BABOONS ,MlD THE BACKGROUND: GENERAL IN'l'RODUCTION"

ABSTRACT 3

1.1 INTRODUCTION 4

1.2 THE MANAGEMENT OF RELATIONSHIPS: THE BACKGROUND

AND THE PROBLEM 6

1.3 THE STUDY OF RELATIONSHIPS IN ANI~~LS: A

CONCEPTUAL FR~~EWORK 10

1.4 MALE BABOONS: SUITABLE SUBJECTS FOR THIS STUDY 12

1.5 A MODEL OF RELATIONSHIP MANAGEMENT FOR ¥~LE BABOONS 17

1.6 OUTLINE OF DISSERTATION 20

CILA..pTER TWO
"THE CONCEPT }-~~D DEFINITION OF DOMINANCE IN ANIMAL BEfI~VIOUR"

l>.BSTRACT 23

2.1 INTRODUCTION 24

2.2 CATEGORIES OF DOMINANCE DEFINITIONS 25

2.3 COMMON DEFINITIONS OF DOMINANCE 27

2.4 GENERAL ATTRIBUTES OF DOMINANCE DEFINITIONS 32

2.5 DOMINANCE AND HERITABILITY 35

2.6 DESCRIPTIVE, PREDICTIVE AND EXPLANATORY VALUE OF

DOMINl>.NCE 36

2.7 THE (IN)ADEQUACY OF DOMINANCE DEFINITIONS 41

2.8 THE FEATURE RECOGNITION CRITERION AND DOMINANCE

RELATIONSHIPS 45

2.9 INDIVIDUALS, SEX, GROUPS, OR SPECIES: THE UNITS

OF DOMINANCE 47

2.10 THE ESSENCE OF DOMINANCE: SYNTHESIS AND DEFINITION 49

1
 PAGE
C!L~PTER THREE
"METHODS"

ABSTRP.CT 53
3.1 STUDY SITE 54

3.1.1 Mikumi National Park 54

3.1. 2 Gombe National Park 55

3.2 STUDY SUBJECTS: THE BABOONS 56

3.2.1 Yellmv baboons 56

Background of the study troops 56

3.2.2 Olive baboons 57

History of the Beach Troop 58

3.3 OBSERVATION PERIODS 59

3.4 DEFINITIONS OF AGE AND SEX CLASSES 61

3.5 COMPOSITION OF STUDY POPULATIONS AND TROOPS 62

3.6 THE MALE BABOONS OF THIS STUDY 64

Residence status and list of individual males 67

3.7 DATA COLLECTION 70

3.8 DEFINITIONS OF BROAD BEHAVIOUR CATEGORIES 72

3.9 NOTES ON STATISTICS 73

3.10 SUM¥~RY OF DATA AND OBSERVATION TIME 74

3.11 ETHICAL CONSIDERATIONS AND BIAS FROM HUMAN INTERFERENCE 75

CfU..PTER FOLTR
"BEP.AVIOURA..L COR..llliU.TES OF SPATIAL DOMINA.~CE STATUS"

ABSTRACT 77

INTRODUCTION 78

METHODS 80

Definitions of behaviours relevant to this chapter 80

RESULTS 83

4.1 Dominance status assignment from spatial outcome of

interactions 83

4.1.1 Definition of procedure and outcome of interactions 83

4.1.2 Assignment of dominance status in males dyads 84

4.1.3 Validation of "unresolved" relationships 87

4.2 Asymmetries in rates of various interaction types 88

4.2.1 General patterns 88

4.2.2 Reversals in resolved dominance dyads 89

4.2.3 Frequencies of aggression, gentle contact and supplants 90

4.3 Asymmetries in socio-sexual and other behaviours 91

4.3.1 Levels of statistical independence 91

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 TABLE OF CONTENTS

PAGE

4.3.2 Results 93

4.4 Mounting behaviour between male baboons 97

4.5 Spatial and ethological criteria in unresolved dyads 99

DISCUSSION 102

4.6 Interaction frequencies in male dominance dyads 102

4.7 Unidirectional behaviours in Mikumi and Gombe baboons 104

4.8 Differences between Mikumi and Gombe baboons 107

4.9 Mounting behaviour 110

4.10 Methodological implications 112

4.11 Behaviour and dominance: a correlation 114

CONCLUSION 118

Cp~1t...pTER FIVE
"SOCIAL DETER..JHN]>~"!TS OF INTE..~CTION R..~TESn

ABSTRACT 119

INTRODUCTION 120

METHODS 126

RESULTS 127

5.1 Interaction rates and relative rank of recipient 127

5.2 Selectivity among potential interactants 127

5.3 The subordinate's perspective towards selectivity

by dominants 130

5.4 Interaction rates and residence status 131

5.5 Interaction rates and number of males in the troop 132

5.5.1 Absolute interaction rates 134

5.5.2 Per capita interaction rates 135

5.6 Changes in interaction rates within dyads 136

5.7 Relationship between individual rates of

interactions initiated and received 140

5.8 Interaction rates and changes in dominance status 142

5.9 Resource vs. non-resource related interactions 146

5.9.1 Proportion and qualities of non-resource interactions 146

5.9.2 Frequencies of resource and non-resource

related interactions 149

Dyadic variation in hourly rates 150

Dyadic variation in proportions 151

DISCUSSION 152

iii
 PAGE
CK~PTER SIX

"l>..N INDEX OF SPATIAL INTOLEP..l>..NCE TO MONITOR THE

DYNAMICS OF SOCIl>..L RELATIONSHIPS"

ABSTR.1..CT 163

INTRODUCTION 164

METHODS 167

RESULTS 169

6.1 General spatial pattern of approach and responses 169

6.2 Different approaches and the likelihood of

eliciting a retreat 172

6.3 The Intolerance Index 175

6.4 Properties of the scale of spatial intolerance 177

6.5 Differences in proximity intolerance between

dominant and subordinate males 180

6.6 Intolerance in resolved and unresolved dominance dyads 182

6.7 Association between intolerance by the superior

and intolerance by the inferior male in dyads 186

6.8 Intra-troop variation between dyads in mean proximity

intolerance 188

6.9 Mean intolerance index of three alpha-beta dyads 189

6.10 Changes within dyads in proximity intolerance

over time 191

6.10.1 Overall variation within dyads in proximity

intolerance over time 192

6.10.2 Changes in proximity intolerance during the

initial phase of relationships 194

6.10.3 Magnitude of changes in proximity intolerance by

subordinate males among dyadic relationships in Mikumi 196

6.10.4 Proximity intolerance by one individual

towards several other males and troop

synchronization of changes over time 197

6.11 variation between dyads in mean intolerance

indices and differences in dominance rank 198

DISCUSSION 202

6.12 On the origin of the intolerance index: a synopsis 202

6.13 The economics of approaches and their responses 203

6.14 Validity and level of resolution of the

intolerance index 204

6.15 Intolerance index and dynamic relationships 206

6.16 Proximity intolerance and rank distance 209

6.17 Other applications for the intolerance index 210

CONCLUSION 211

iv
 TABLE OF CONTENTS

PAGE
CF..APTER SEVEN
"SPATIl<..L INTOLERA.~CE, INTERACTION RATES A.~D INTIMIDATION"

ABSTR~CT 213
INTRODUCTION 214
METHODS 215
RESULTS 216
7.1 Proximity intolerance and interaction rates 216
7.2 Interactions initiated by the subordinate and the
subordinate's intolerance of the dominant 219
7.3 Change in dominance status, proximity intolerance and
interaction rates 220
DISCUSSION 224
7.4 Evidence of an effect between interaction rates
and intolerance 224
7.5 Interaction rates and spatial
intolerance: determining causality 225
7.6 The possible influence of interactions with third
parties on the spatial relationship between two males 226
7.7 The case for intimidation between male baboons 226
7.8 The function of intimidation derived
from human empathy 227
7.9 Further evidence against a model of dominance based on
variation in competitive ability 230
CONCLUSION 230

CF..Jt.J>TER EIGHT
"GENTLE CONTACT, AGGRESSION, SUPPLA.~TS AND Ml>..LE REL..~TIONSHIPS"

ABSTRb.CT 233
INTRODUCTION 234
METHODS 237
RESULTS 238
8.1 Overall proportions of different interaction kinds 238
8.1.1 Proportions of gentle contact, aggression
and supplants 238
8.1. 2 composition of gentle-contact interactions 239
8.1. 3 composition of aggressive interactions 240
8.1. 4 composition of supplants over resources 240
8.2 The resource context of interactions 242

v
 PAGE

8.3 Correlations between rates of aggression, gentle

contact and other interactions 244

Intertroop differences 245

8.4 Discriminating between functions of gentle contact:

appeasement or antagonism 247

8.5 Submissive behaviour during

gentle-contact interactions 247

8.6 Gentle-contact and aggression among

resolved and unresolved dominance dyads 251

8.7 Spatial intolerance and frequencies of gentle

contact and aggressive interactions: a correlation

analysis across dyads 254

Null hypothesis and variables analysed 254

8.7.1 General results 256

8.7.2 Specific results 258

8.8 Spatial intolerance vs. rank distance 262

DISCUSSION 263

8.9 Are gentle-contact interactions affiliative? 264

8.10 Social interactions and immediate resource acquisition 268

8.11 Relationship management through

interactions: the evidence 269

8.12 Constraints on rank distance as a parameter of

relationships 273

8.13 Differences between Troop 2 of yellow baboons and

Troop 5 of olive baboons 274

8.14 Psychological warfare between males: an analogy 275

8.15 Conclusions 276

CP..APTER NINE
"COALITIONS A.~D ~~..LE RELATIONSHIPS"

ABSTRz>'CT 277

INTRODUCTION 278

METHODS AND DEFINITIONS 280

RESULTS 281

I. General patterns 281

9.1 Broad categories of coalitions involving males 281

9.2 Request for support as a trigger of coalitions 283

9.3 Partners and targets of parallel coalitions 284

9.4 Partners and targets of interference coalitions 285

vi
 TABLE OF CONTENTS

PAGE

9.5 Adult males as partners during interference

and parallel coalitions 287

9.6 Concluding remarks 288

II. Coalitions of two or more males 289

9.7 Number of participants and definition of roles 289

9.8 Outcome of coalition attacks 290

9.9 Aggression levels during coalition attacks 290

9.10 Dominance relationships and risky interventions 292

9.11 Resource context of coalitions 295

9.12 Take-overs of oestrous females through coalitions 297

9.13 Defense of consortship through coalition 302

9.14 Characterization of alliances 303

9.15 Affiliative relationships between allies? 307

9.16 Reciprocity of support given in coalitions 309

9.17 Emigration and rate of being the target of coalitions 312

DISCUSSION 316

9.18 Costs of participation in coalitions 317

9.19 Resource context of coalitions 318

9.20 Motive for helping 1: reciprocation in the future 320

9.21 Motive for helping 2: skill acquisition 321

9.22 Motive for helping 3: benefits from

relationship management 322

9.22.1 Fostering alliances and benefits from reliability 322

9.22.2 Dependent rank among male baboons: an old

concept revisited 324

9.23 Dyadic relationships and partners in

coalitions: a link? 325

9.24 Conclusions 328

CI-LAJ>TER TEN
"PATTERN, CONTEXT l>..ND COSTS OF INJURIES IN MALE BABOONS"

ABSTR~CT 329

INTRODUCTION 330

METHODS 331

RESULTS AND DISCUSSION 333

10.1 Fights and the baboon's weaponry 333

vii
 PAGE

10.2 Types of injuries 335

10.3 Contexts of injury and contexts of no injury 336

10.4 Frequency of injury 338

10.5 Bodily location of injuries 341

10.6 Healing time 344

10.7 Number of wounds per fight 345

10.8 Determinants of wounds per fight: oestrous females? 347

10.9 Determinants of wounds per fight: male immigration? 348

10.9.1 Relationship between wounds per fight and days

since immigration 348

10.9.2 Probability of fights of any number of wounds

occurring soon after male immigrations 349

10.9.3 Immigration context of fights resulting in high

numbers of wounds 349

10.10 Costs of injury 351

Reduced competitive ability and drop in rank 352

Feeding impairment 355

Lethal injuries sustained in fights 355

10.11 Concluding remarks 356

CP~~PTER ELEVEN
"PSYCHOLOGICAL WARF]'~~E A.~D THE fI.ANAGEMENT OF RELATIONSHIPS
BETWEEN fI.ALE BABOONS: GENERl·J., DISCUSSION"

ABSTRACT 359

INTRODUCTION 360

11.1 SEPAP~TING FACTS FROM SPECULATION: AN

OVERVIEW OF RESULTS 361

11.2 THE MECHANISM OF INTIMIDATION 365

11.3 GAME THEORY AND BABOON INTEP~CTIONS 369

11.3.1 Introduction 369

11.3.2 The first encounter between two males 371

11.3.3 The second and subsequent interactions in

dominance dyads 374

11.3.4 Bluffing and honesty in male baboon interactions 380

11.3.5 Implications for the theory of aggressive competition 386

11.3.6 Conclusion 387

11.4 THE FUNCTION. OF RELATIONSHIP MANAGEMENT

IN ~~LE BABOONS 388

11.5 COGNITIVE ISSUES AND RELATIONSHIP MANAGEMENT 392

11.6 THE CASE FOR PSYCHOLOGICAL WARFARE IN MALE BABOONS 398

viii
 TABLE OF CONTENTS

PAGE

11.7 MALE BABOON RELATIONSHIPS: THE FINAL PICTURE 402

11.7.1 General features 402

11.7.2 Male baboons and their relationships: a synopsis 405

11.7.3 Some implications and suggestions for

complementary research 407

11.8 RELATIONSHIP ~~.NAGEMENT: BABOONS AND BEYOND 408

11.8.1 Bridging the gap between species 408

11.8.2 Dominance as a low-risk state of imprecise knowledge 410

11.8.3 Dominance vs. competitive ability: an

unresolved paradox 411

11.8.4 A deterministic view on dyadic aggression 412

11.8.5 Support for the Machiavellian

Intelligence Hypothesis? 413

11.8.6 Baboons, human empathy and generating hypotheses 414

ACKNOWLEDG&~ENTS 417

APPENDIX 1. SOME DIFFERENCES BETWEEN YELLOW

AND OLIVE BABOONS 419

APPENDIX 2. FREQUENCIES OF DIFFERENT AGGRESSION LEVELS 431

APPENDIX 3. A CASE OF TACTICAL DECEPTION IN MIKUMI BABOONS? 435

LIT~~~~~~E CITED 437

REh~TIONSHIPS, BABOONS P~D THE BACKGROUND: GENER~ INTRODUCTION

ABSTR~CT

It is of paramount importance to understand social relationships

between animals in order to understand how gregarious behaviour
evolved and hOvJ animals take advantage of a social setting to
pursue their individual goals. Active management of relationships
can be expected, given the competiti'Ie advantage that it might
confer. Previous studies suggest that long-term social motives can
influence patterns of interactions between animals that live in
stable groups. Few studies, however, have specifically addressed
the management function of mundane interactions in competi ti 'Ie
relationships. This study concerns the social goals, sometimes
Machiavellian, that motivate male baboons (Papio cynocephalus) when
they interact with each other. Male baboons are suitable subjects
for the topic of this dissertation for several reasons. They can
be studied in their natural habitat with relative ease. Baboons
live in groups containing several unrelated adult males. Males
transfer singly between troops, which makes their relationships in
any given troop relatively short-lasting. Consequently, the
monitoring of male relationships can include their genesis,
development and termination. The conceptual framework for social
relationships in animals and people developed by R.Hinde (1976, Man
11: 1-17) is used in this investigation. A central, original
feature of this study is the quantification of the effect of
management through interactions on the quality of relationships.
The results are relevant to our understanding of the role of
interactions in animal societies, as well as of the subtle
mechanisms that underlie competition.

3
 THE CONCEPT ~~~D DEFINITION OF DOMIN~~~CE IN ~~~IP~~ BEP~~VIOUR

ABSTR~CT

The concept of dominance has contributed greatly to our

understanding of social structure in animals. Over the past three
decades, however, a variety of concepts and definitions of
dominance have been introduced, leading to an ongoing debate about
the usefulness and meaning of the concept. Criticisms aimed at one
definition of dominance do not necessarily apply to other
definitions. Existing definitions can be structural or functional,
refer to roles or to agonistic behaviour, regard dominance as a
property of individuals or as an attribute of dyadic encounters,
concentrate on aggression or on the lack of it, and be based either
on theoretical constructs or on observable behaviour. Thirteen
definitions of dominance are reviewed, and their usefulness
assessed with respect to their descriptive value. The predictive
and explanatory values of definitions are specific to the questions
asked in each particular study and are not considered as criteria
to judge the usefulness of the dominance concept. By virtue of its
high descriptive value, the original definition of dominance by
Schjelderupp-Ebbe (1922, Z.Psychol. 88: 226-252) emerged as the
basis to formulate a structural definition with wide applicability
and which reflects the essence of the concept: Dominance is an
attribute of the pattern of repeated, agonistic interactions
between two individuals, characterized by a consistent outcome in
favour of the same dyad member and a default yielding response of
its opponent rather than escalation. The status of the consistent
winner is dominant and that of the loser subordinate. Dominance
status refers to dyads while dominance rank, high or low, refers
to the position in a hierarchy and, thus, depends on group
composition. Dominance is a relative measure and not an absolute
property of individuals. The discussion includes reference to the
heri tabili ty of dominance, application of dominance to groups
rather than individuals, and the role of individual recognition and
memory during agonistic encounters.

23
 METHODS

J>J3STHACT

The behaviour of males in five habituated troops of baboons

was studied in their natural environment. Four of these troops
were observed in Mikumi National Park, Tanzania. Life history
information on the study individuals in two of these troops
was made available to this investigator from the Mikumi Long
Term Baboon Project, which has operated since 1975. The yellow
baboons in Mikumi National Park were observed in two periods
representing a total of 16 months, between May 1990 and
November 1991. The habitat in Mikumi National Park is
characterized by Miombo woodland on the hills, and open
woodland and grassland in the valley.
In addition to the Mikumi troops, one troop of
habituated olive baboons was observed for six weeks in Gombe
National Park, in western Tanzania. Observations in Gombe took
place in September and October 1991. Rainfall is much higher
in Gombe than in Mikumi. Common vegetation zones in the range
of the Gombe study troop was the evergreen and the deciduous
forest as well as the beach on the lake shore.
Baboons \vere follmved on foot avoiding interference \.Ji th
the behaviour of the animals. The observations were logged on
check-sheets by two observers. Social interactions between
males were recorded ad libitum and in focal animal samples
of 15 min duration. The baboons were well habituated to human
observers and could be observed at close range. The data set
collected corresponds to 1,413 hours of baboon observations,
of which 532.5 hours were focal observation time. The main
study troop accounted for 52% of the observation hours. Over
2,100 interactions between adult males were recorded.

53
 BEH..~VImJR:lili COR..llliL~TES OF SPATI.A..L DOMINANCE STATUS

.AJ3STRACT

Patterns of approach and retreats are the spatial behaviour which

was chosen in this study to characterize the relationships between
male baboons. The loser of an interaction was defined as the
individual which retreated. Following the definition of dominance
proposed in Chapter 2, dominance status between brJO males \..;ras
assigned only if there was a statistically significant asymmetry in
the series of outcomes of their agonistic interactions. Assignment
of dominance status from spatial behaviour is biologically
meaningful. The majority of male dyads had a resolved relationship,
in which most interactions were initiated and won by the dominant.
several dyads of Gombe baboons had unresolved relationships. Some
dyads with few interactions were classified as unknown relationships.
A subordinate sometimes elicited a retreat from a dominant, but such
reversals represented on average less than 10% of all interactions
in resolved dominance dyads. About a third of interactions received
by the subordinate resulted in the loss of a resource as it
retreated, whereas the dominant rarely lost a resource upon being
approached by the subordinate. Interactions initiated by the
subordinate included a greater proportion of gentle contact
interactions than interactions initiated by the dominant. The spatial
criterion was further validated by its overall agreement with
ethological criteria which include the direction of aggressive and
submissive behaviour in the definition of status. In Mikumi and Gombe
baboons the strongest behavioural indicator of subordinate status was
the grimace and its associated call, the gecker, which have been
considered submissive behaviour in previous studies of
cercopithecines. The Mikumi and Gombe baboons differed, however, with
respect to the pattern of hindquarter presentations and raised tail,
among other behaviours. The variety of behaviours associated with
dominance status show that dyadic dominance is an important parameter
to characterize the relationships between adult male baboons.

77
 ~OCI~~, DETE.~~INANTS OF INTERACTION RATES
ABSTRA,.CT
The working hypothesis was proposed that animals manage their
relationships principally through variation in the frequency of social
interactions. It was predicted that the rate of interactions between
b·JO animals vJOuld be, at least partly, determined by qualities of
their relationship and that, therefore, animals should choose
selectively the target and frequency of their interactions. Several
specific predictions of the model of relationship management \·..ere
confirmed among the study baboons. Males initiated most interactions
towards subordinate males close in rank, as would be expected from
efforts to reassure their dominance status. Selectivi ty was also
influenced by residence status, whereby residents tended to
concentrate their interaction on newcomers, and newcomers tended to
allocate (high rates of) interactions evenly among the residents.
Frequent interactions between residents and newcomers could be a means
of gathering information about the quality of competitors, as well as
cementing dominance relationships in their early stages. Non-resource
interactions, representing about half of all interactions betv..een
males, were used selectively towards newcomers by the alpha male of
Troop 2. Absolute rates of interactions initiated and received were
influenced by dominance rank and by the number of males in the troop.
Lov..-ranking males received more and initiated fewer interactions than
higher-ranking ones. For any given male, the rate of initiated
interactions increased with the number of subordinates, and thus with
the number of relationships. The rate of interactions received
increased \'lith the number of dominants, hence illustrating a cost
factor for males occupying low ranks. Within male dyads, however, the
rates of interactions received and initiated were not related in a
simple 'i.·.. ay. The role of social factors ln determining rates of
interactions \·;as particularly obvious in bvo cases of dominance
reversals. Several weeks before the change, the dominant male
increased considerably the frequency of interactions tm·;ards the
subordinate, perhaps in an attempt to reaffirm his status. The changes
in status were abrupt, and were associated with drastic changes in the
pattern of interactions between both males. As an alternative to the
model of relationship management, a more parsimonious model of
immediate circumstances could explain some of the patterns found.
Overall, however, the bulk of evidence from this and other studies
suggests that long-term social goals can determine interaction
frequencies in animals.

119

 ~~~ INDEX OF SPATIAL INTOLE~~CE TO MONITOR THE DYNp~ICS OF

OF SOCIAL RELATIONSHIPS
ABSTR~CT

One of the proposed functions of social interactions is to reinforce

existing relationships, either by strengthening affiliative bonds
or by reassuring dominance status. The effect of interactions on the
quality of relationships, however, has not been quantified because
of lack of an independent measure to characterize relationships.
In this chapter, the spatial information contained in approach­
response interactions between male baboons was used to develop such
a measure: the spatial intolerance index. The index indicates for
each male how intolerant it is of the proximity of another male. It
is a measure of a male's reluctance to retreat upon being
approached. The intolerance scale combines fast, slow, distant and
close approaches, as well as the possible responses, including
counter-approaches, ignoring, and retreats of various types. Each
interaction between males is given its corresponding intolerance
index. The mean of all indices recorded is calculated within dyads
separately for each male as the recipient of approaches by the
other. As expected from a spatial model of dominance, the proximity
intolerance of subordinates by dominants was lower than vice versa.
Therefore, the index is biologically meaningful. It has a useful
level of resolution. Significant variation in spatial intolerance
was detected both between dyads and within dyads over time. Male
relationships were revealed by the index as dynamic, even when
changes in dominance status did not occur. The subordinates'
intolerance of dominants, in dyads involving newcomer males,
decreased during the first 40 days of the relationship, and
increased thereafter to the initial levels. Some dyads showed long­
term oscillations in intolerance, whereas in one case a steady
decrease in intolerance by the subordinate was documented. The mean
intolerance by the dominant was v.leakly, and not significantly
correlated with the subordinate's intolerance. In general, each
subordinate male showed similar intolerance levels towards all his
dominants. Differences between subordinates in their intolerance of
high rankers, however, could be significant. Rank distance between
males does not affect intolerance levels in a simple way for males
seem to simply discriminate between other males above and below
their own rank. The pattern of approaches and responses between male
baboons is at least partly determined by social qualities of each
relationship rather than merely by immediate ecological
circumstances. The spatial intolerance index can, among other
things, be used as a lens to study how relationships are managed.

163

 SPATI~~ INTOLEP~~~CE, INTEP~CTION P~TES ~~D INTIMIDATION

p..BSTRACT

Interaction rates follow patterns consistent with the model of

relationship management and were proposed in Chapter 5 as the
mechanism to achieve long-term social goals. This chapter tests the
prediction that interaction rates should affect the quality of
relationships. In an analysis within dyads, the relationship between
interaction rates and spatial intolerance is investigated. The non­
random pattern of changes in intolerance within male dyads over time,
is associated with variations in the rate of interactions initiated
by the dominant, at least in some cases. In two dyads involving the
alpha male, the level of spatial intolerance by subordinates vJaS
significantly, posi ti vely correlated vIi th the rate of approaches
received from the dominant. This represents original, quantitative
evidence of an association between the rate of interactions initiated
by an animal and a change in its relationship with the recipient. The
intolerance index visualized the effect of persistent use of
displacements on the attitude of a subordinate towards a higher
ranking male. This process can be referred to as intimidation. The
separate relationships between one subordinate and b ..J O dominants
pointed to the possibility that one of the dominant males, the
"harasser", was shaping the subordinate's level of intolerance shown
towards all dominants. Increased interaction rates by the dominant
male preceded changes in dominance status in two additional dyads.
These can be interpreted as attempts to reassert dominance status. In
one dyad, the low level of intolerance shown by the subordinate,
before the change In status, approached the bold behaviour shown
generally by dominants. Conversely, in the other dyad the level of
intolerance by the subordinate increased following the sharp increase
in received interactions, possibly showing that intimidation was
effective. The change in status in that dyad was associated with a
sUbsequent injury of unknown origin sustained by the dominant. It is
argued that the main function of gradual intimidation by dominants is
to facilitate the future access to mating opportunities through low­
risk appropriation of oestrous females and low-risk defence of
consortships from coalitions.

213
 GENTLE CONTACT, AGGRESSION, SUPPLANTS AND MALE RELATIONSHIPS

ABSTRACT

In this chapter, the function of social interactions was

investigated following a working hypothesis of relationship
management. Firstly, it was established that gentle-contact
interactions are not affiliative in nature and this lead to the
conclusion that affiliative interactions between adult male baboons
are very rare if not completely absent. Although overt aggression
is generally absent from gentle-contact interactions between males,
these interactions are still best regarded as a mild form of
antagonism. The rare occurrence of resource appropriation resulting
from gentle contact or aggression suggests that immediate resource
competition is unlikely to be the main motive of these and other
approach-retreat interactions, which are the bulk of interactions
between males. Instead, it is proposed that the achievement of
long-term social goals is the main function of these interactions,
with some reservation about the role of aggression in this process.
The role of interactions in the long-term servicing of
relationships was supported by several statistically significant
correlations between frequencies of dyadic interactions and the
spatial intolerance relationship between males of Troop 2 of yellow
baboons. The validity of these results across baboon subspecies is
uncertain, despite similarities in the dyadic proportions and rates
of different interaction types in yellow and olive baboons. The
spatial intolerance index developed in Chapter 6 was instrumental
in the identification of the above associations. Rank distance
failed to detect any of these associations, partly as a consequence
of dealing with a small group of individuals, but also due to
weaknesses inherent in the use of rank distance to characterize
relationships. In conclusion, males appear to be involved in a
constant psychological war based on persistent, overtly peaceful
approaches.

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 COALITIONS AND MALE RELATIONSHIPS

ABSTRACT

Coalitions between male baboons have been principally regarded as

a means to gain access to oestrous females consorted by higher
ranking males. Most coalitions, however, are not in the context of
immediate mating competition and in many cases occur without
apparent reason. This chapter focuses on the possibility that the
function of coalitions between male baboons lies in part in the
management of their relationships. Analyses include a detailed
description of the structure of coalitions in the study troops,
aggression levels, resource context, alliances, consort take-overs,
reciprocity of support and the effect of coalition rates on the
onset of emigrations. Support for the management of relationships
hypothesis comes from the fact that a large proportion of
coalitions do not take place in the context of tangible resources
and therefore are likely to involve social goals. These social
goals must be important to both partners of coalitions since they
must override the evidently high risk involved in these highly
aggressive interactions. Some likely motives for participation in
coalitions are reciprocation (when access to oestrous females or
protection from an attack are at stake), acquisition of social
skills required in polyadic interactions, identification of allies
and fostering of the alliance, as well as advertisement of
dependent rank. These motives are not mutually exclusive. Partners
in a coalition may differ in their motives to participate. Dyadic
relationships and participation in coalitions are separate worlds
which find a link via dependent rank: subordinates may manage their
dyadic relationships with dominants by participating in coalitions
against them. Dyadic social interactions between males are
seemingly not geared towards the potential for regular
participation in coalitions as partners. One additional dimension
of the instrumental use of coalitions can be the eviction of
immigrant males by several residents, a case of cooperation.

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CHA.PTER. TEN

PATTERN, CONTEXT AND COSTS OF INJURIES IN MALE BABOONS

ABSTRACT
The pattern, context and costs of injury in male baboons was
investigated to establish their possible role as determinants of
peaceful conflict resolution and psychological warfare. Most of the
findings reported here for male baboons apply also to males of other
multi-male primate species. Patterns of injury and fights are mainly
a consequence of the nature of the baboon weaponry. The upper
canines are the main weapons used in fights between male baboons,
and are capable of inflicting severe and sometimes lethal injury.
During contact fights both opponents risk injury regardless of
differences in competitive ability. The vast majority of wounds in
adult baboons result from intraspecific aggression. Wounds are more
common in males than females. In males they concentrate on anterior
parts of the body as expected from face-to-face combat, and most
wounds are inflicted on the right side. Over 80% of the males were
injured at some point during the study. The individual rate of
injury from fights with other males is once every 1.5 months. The
number of wounds sustained in each fight was not related in a simple
way to availability of oestrous females or recent immigration
events. The four fights yielding the highest number of injuries,
however, involved recent immigrations or attempts to immigrate by
young males in their prime. Such challenges to the resident alpha
male and fights over oestrous females probably constitute the main
contexts leading to injury. Encounters over preferred, rare foods
and some fights unrelated to tangible resources also resulted in
injury. The costs of injury to male baboons can be substantial.
Physical impairments resulting from wounds sustained in fights can
constrain feeding efficiency and ease of access to resting sites as
well as to safe retreats. More importantly, injury can reduce the
competitive ability of males, jeopardize their mating success and
even cause a drop in dominance rank. Male baboons respond
behaviourally to such social changes by avoiding further
confrontations, retreating to the periphery of the troop or
emigrating temporarily. The most severe fights between males can
result in death of a combatant. Lethal fights are probably not
uncommon in baboons and tend to be associated with coalition
attacks. The potentially high costs of fights between males are the
premise for psychological warfare through subtle approach-retreat
interaction (Chapter 11).

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 PSYCHOLOGICAL WARFARE AND THE MANAGEMENT OF RELATIONSHIPS
BETWEEN MALE BABOONS: GENERAL DISCUSSION

ABSTRACT
This chapter amplifies the interpretation of the results of this study
by addressing complementary aspects such as the underlying mechanisms,
evolutionary and cognitive issues, as well as by framing the findings
in a wider context. The model of relationship management was found to
be plausible from the point of view of the proximate behavioural and
physiological mechanisms involved in intimidation. Evolutionary
considerations, based on the theory of games, also allowed the
conclusion that the process of social management is plausible in that
the behaviour patterns observed can, in principle, evolve according to
the functions proposed. The facts that competitive ability in male
baboons is largely concealed and that the costs of escalation are high
provide an adequate scenario for deceitful behaviour to be stabilized
by natural selection. The interactions between males are likely to have
elements of blUffing as well as honest advertising of their competitive
ability, and in addition can serve the assessment of the opponent's
capabilities. The function proposed for such exchange of information,
in which both the dominant and subordinate males are actively involved,
is to minimize ultimately the risk of escalation in subsequent
encounters over mating opportunities. The effects of chance and
mistakes in assessment during initial stages of the relationship are
believed to result in dominance relationships which do not necessarily
conform to a ranking based on competitive ability. Imperfect assessment
is the basis for the dynamic processes which prevail in male
relationships. Males live in a state of constant dyadic competition
which is best described as "psychological warfare". In a climate of
tension, in which the risk of escalation is latent in each interaction,
most encounters between males consist of subtle approach-response
events in which social rather than immediate tangible goals are at
stake. The current state of knowledge about cognition in monkeys
suggests that male baboons are unlikely to show intentional behaviour
or to attribute mental states to other males, but that they are able
to feel the pain, fear and anxiety associated with their warfare in a
way similar to humans. This discussion addresses the applicability of
the results to other species, as well as the implications to our
understanding of dominance, aggression and competition in animals.
Human empathy and introspection, the seeds for the working hypothesis
of this study, are considered powerful tools for unraveling the
complexity of primate behaviour.

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