THE

CONCEPT

AND

DEFINITION

ANIMAL

OF DOMINANCE

IN

BEHAVIOUR
by

CARLOS DREWS1)
of
Zoology, Cambridge University, Downing Street, Cambridge CB2 3EJ,
(Department
U.K.)
(With 1 Figure)
(Acc.7-VII-1993)

Abstract
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 necessarilly 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 heritability of dominance,
application of dominance to groups rather than individuals, and the role of individual
recognition and memory during agonistic encounters.
Introduction
Dominance

is a keyword in most studies of social behaviour in gregarious

animals and has contributed
of social strucgreatly to our understanding
ture. Tables listing dominance
studies in a diversity of animal taxa are
G.W. NORTON,
and P.C. LEEfor valuable discussion.
1) I thank S.K. ELTRINGHAM,

284
provided by DEWSBURY (1982) and GAUTHREAUX (1978). Despite the fact
that dominance
is such an important
and widely used concept, there is
still no agreement
its meaning.
regarding
SCHJELDERUPP-EBBE (1922)
introduced
the concept of the peck-order,
later to be called dominance,
into the behavioural
sciences. He vividly described his observations
on
the peck-order
and the
of domestic fowl, the dynamics of the hierarchy,
determinants

of rank.

types of domifound in animals. HAND (1986) provided

an ample
nance hierarchies
discussion of the different categories found among dominance
relationdefined the terms commonly
used in studies of
ships and rigorously
dominance.

WILSON (1975) summarized

different

Other

can be found
conceptual discussions about dominance
in VAN KREVELD (1970), FEDIGAN (1982) and DUNBAR (1988).
A lively debate has underlined
in behavthe application
of dominance
ioural studies over the past three decades (e.g. BERNSTEIN, 1981). GARTLAN
of dominance
because these
(1968) criticized the different interpretations
often

led to ambiguity.
He further criticized
the fact that the term
is often used without
a definition
or else is arbitrarily
redefined to fit the findings. ROWELL (1974) pointed out that the definition of dominance
is elusive. Investigators
are far from a consensus on
dominance

how to define

and measure

dominance

(BERNSTEIN, 1981), leading to

and evolution. BERNSTEIN (1981)

disparate opinions regarding its function

concentrated
on primates but his review of the problems associated with
the concept of dominance
has general application.
His article is accomfrom
panied by comments
overview of the dominance
One

of the central

various
debate

and thus provides an

investigators
and lack of concensus at that time.

of the dominance

problems

debate

has been

that

critics have not always explicitly specified which definitions they advocate
they criticize. This has kept the debate going since the criticisms may have been justified with respect to a particular
definition but
and which

dominance

In this paper, an overview of the diversity of definitions of

found in the animal behaviour literature is presented and the

usefulness

of each

not to others.

understanding
definitions
and

definition

of social

assessed

structure

with respect to furthering

our
and behaviour.
Elimination
of some

denominator
of others lead to a
finding the common
in
the
essence
of
dominance
behavioural
studies.
illustrating

synthesis
This synthesis
the concluding

is condensed
section.

in the definition

of dominance

proposed

in

285
of dominance

Categories

definitions

The

terms winner, loser, submissive, subordination

and aggression are
of
the
the
dominance
specialized vocabulary
surrounding
concept.
part
These terms are defined and used here as follows:
Winner:the contestant that expresses consummatory behaviour according to its initial goal
1981; HAND,1986). Winner/loser refers to the outcome of one single contest.
(BAENNINGER,
Submissive:submissiveness is an active response to an aggressive action by another individual (HANBY,1976) in which yielding or surrendering are displays given by the loser (HAND,
1986). Submissive behaviour refers to single contests.
Subordination:to lose consistently certain conflict encounters (HAND,1986). Subordination
refers to a pattern based on several contests.
Aggression:actual attacks, threats of attack or encroachments; signals which suggest that
attack may occur can be called aggressive (HAND,1986).
into those based
can be separated
based on theoretical constructs

Dominance

definitions

observations

and definitions

HINDE & DATTA, 1981 ). These have been referred

and "theory-language"
definitions
respectively
two approaches
( 1981 ). GAGE ( 1981 ) distinguishes

on empirical
(GAGE, 1981,

to as "data-language"
by HINDE & DATTA

to the study of domiin theory-language

nance : in the first, the investigator
formulates
the
conditions which must be met by a useful application
of the term domi-

nance,

a testable

then derives

hypothesis and lets the results of empirical

or else lead to its rejection. In
of dominance
the investigator
assumes the existence of domi-

the definition

inquiry support
the second approach,

the
asserts an operational
definition, and proceeds to demonstrate
This latter approach
is similar to the
utility of the term and definition.
definitions
mentioned
HINDE
DATTA
&
by
(1981), in which
data-language

nance,

a particular behavioural
action". More broadly,
set of observable
empirical
functional
describe

interaction
dominance

interactions.

observations

can

is given the name "dominance

interis used to describe a particular kind or
The dominance
definitions
based on

in turn

be subdivided

into

structural

and

ones (BERNSTEIN, 1981). Structural

dominance
definitions
the pattern of an observable
or
set
of
interactions
without
type
to their function, whereas functional definitions describe domi-

referring
nance from the point of view of its apparent function.
The distinction
between definitions
observations
based on empirical
and theoretical
as well as between structural
and functional
postulates,
definitions

is used in this discussion

to broadly

characterize

the definitions

286
of dominance

presented below. A further distinction between definitions

is whether they describe agonistic or non-agonistic
dominance
relationon
the
definitions
of
dominance
based
First,
ships (BERNSTEIN, 1981).
empirical
definitions

observations

are presented.
These are ordered from broad
to more specific ones and, as such, they are not mutually
is given.
based on theoretical
constructs
exclusive. Then, a definition
Some definitions are included which are not explicitly referred to in the
literature,

but which

help in the process

Common

definitions

of dominance

Definitions

on empirical

based

of categorization.

observations.

animals in a role which subjectively

a
represents
with
to
others
are
dominant.
This
definition
is
respect
privileged position
illustrated by WILSON'S (1975) description
of the dominance
hierarchy of
to nurses (dominant).
food transfer from forager bees (subordinates)
There is no overt aggression in the relationship
and the subordinate
bees
1. "Privileged

role":

status
acquire dominant
dominance
definition.

as they

grow

older.

This

is a non-agonistic

definitions
include agonistic behaviour
as part of the
remaining
dominance
(2.-4.) are listed which consider
concept. First, definitions
to be an attribute of the individual.
In subsequent
definitions
dominance
of
is considered
a relative measure,
an attribute
(5.-13.) dominance

The

dyadic

encounters

or relationships

and not a property

of individuals.

in social insects refers

status": reproductive
dominance
2. "Reproductive
in
individual
status such that the reproductive
to differences
reproductive
is the dominant and the non-reproductive
ones are subordinates
(WILSON,
be regarded
as a case of the
"privileged role" definition. The use of overt aggression by the reproductives, however, justifies a separate mention of this definition. When two or
more paper wasp (Polistes sp.) females start a nest together, one becomes
1975).

This

structural

definition

could

the egg layer while the others become workers. The queen establishes her
dominant
position and controls the other wasps by direct aggression
refers to the reproduc(WEST, 1967; EBERHARD, 1969). Here, dominance
tive condition achieved and not to the pattern of aggressive interactions.

287
the investigator

If, however,
aggression,
derived from

bases dominance
dominance

then

reproductive
the "dominance

is aggressiveness"

on the pattern of
definition

definition

(see below).

a dominance

is aggressiveness":

3. "Dominance

status

is a functional

is the set of
hierarchy
a
among
group of animals
(WiLSON, 1975). WILSON (1975) is

relations

sustained

aggressive-submissive
coexist within one territory
concerned
with "...dominance
behaviour

which

mediated
and
by aggression
based
natural
selection
at
the
level
of
the
individual...".
inferentially
upon
The occurrence
of aggressive contests over resources among any social
The expression
group of animals is indicative of dominance
relationships.
of

the

dominance

is

system

based

on

individual

variation

in

aggressiveness.
4. "Dominance

is a trait

that

conveys

rank":

a dominant

individual

wins encounters

with significantly
greater than chance probaa
This occurs because that
bility against
variety of conspecific opponents.
individual
has more of a trait called dominance
or
(or aggressiveness
than
do
the
losers
assertiveness)
(BAENNINGER, 1981).).
routinely

5. "\Vinner

is dominant,
loser is subordinate":
dominance
status is a
for winner and loser used to describe the outcome of any single

synonym
dyadic

encounter,

6. "Successful

regardless

combatant":

of whether
dominance

or not the contest

ranks are established

is escalated.
on the basis

of dyadic contests resembling a competition.

Generally each dyad meets
on one or few occasions in which the decisive contest takes place. CLUTToN-BROCx et al. (1979) established
the dominance
deer on the basis of fighting success in the rutting

rankings of male red

season. Similarly, the

ranks of elephant seals are determined

through the outcome of escalated
fights in the breeding season (LEBoEUF, 1974). The observed pattern of
interactions
resembles HAND'S (1986) aggressive dominance
definition, in
which the winner uses or seems willing to use the most aggression.
Dominance

scores based on overall

fighting success belong to this definition (e.g. BOYD & SILK, 1983, ENS et al., 1990). Another
example are
in
which
the
contest
is
sometimes
decided
on
the basis of
lekking species,
This
acoustic or visual displays and may include escalated fights.
defini-

288
implies that escalation is avoided nor that individuals recognize each other and incorporate
into contest
past agonistic experiences
decisions.

tion neither

is lack

of aggressiveness":
dominance
refers to the
nature of encounters
between animals, in which escalated
"peaceful"
conflicts
are resolved using non-agonistic
do
not
take
and
fights
place
VESSEY (1981) advocates a use of the
assessment or submissive behaviour.
term dominance
that conveys its likely function,
namely control of
7. "Dominance

and minimum

resources
control

the behaviour

individual

energy expenditure
through the use of displays to
of the receiver. Learning from past encounters
and
are not necessary conditions under this definition.

recognition
a feature in the opponent
an individual
which
recognizes
Typically,
of
and
submits
indicates superiority
(e.g. size, weaponry,
quality
display)
of outcomes seems necessary under
without a fight. High predictability
this definition

of dominance.

Dominance

status is assigned

after a single

contest.
8. "Consistent

winner

of agonistic contests":
when
against B then A is dominant

agonistic encounters
This definition is characterized

A consistently
wins
and B subordinate.

in the outcome of a
by the asymmetry
It
series of contests of any type and implies that A and B meet repeatedly.
neither implies that high levels of aggression are avoided nor that individual recognition
and memory of other contests play a role in contest
decisions.
in a given context":
dominance
is a descriptive
win dyadic encounters
in a given
term for individuals
that consistently
is
described
for each
context (HAND, 1986). A dominance
relationship
9. "Consistent

competitive

winner

context.
exhibit

can be pure or mixed, since

Dyadic relationships
different
kinds of relationship
for different conflict

they may
contexts (HAND, 1986). This definition emphasizes
the role of context
the outcome of interactions
(e.g. LEE, 1983).
determining

in

10. "Priority of access to resources":

dominance
is by definition priority
of access to resources (VAN KREVELD, 1970; MORSE, 1974; SYME, 1974;
WAGNER &
WILSON, 1975;
GAUTHREAUX, 1978;
WOLFE,
1984;

289
GAUTHREAUX, 1990). Popp & DEVORE (1979) define
in contests over resources.
nate as the winner/loser

dominant/subordi-

The winner is the

or
the opponent's
displaces
appropriates
resource (e.g. CLUTTON-BROCK et al., 1979). Avoidance of escalation durIt refers to obtaining what
ing contests is not a requisite for dominance.
one wants by winning a fight or without having to fight for it (RALLS,
which

contestant

the

other

central statement of dominance

is that particular
1976). The functional,
in
individuals
social groups have regular priority of access to resources in
situations
competitive
(CLUTTON-BROCK & HARVEY, 1976). Dominance
an asymmetry
in the outcome of such contests. High ranking
individuals
can be identified by observing the outcome of contests over
resources or by identifying the holders of resources directly (e.g. the male
describes

with the largest

11. "Peck-order"
never

or seldom

harem

is by definition

the alpha

male).

after SCHJELDERUPP-EBBE (1922): if A pecks at B and B

then A is dominant
to B and B is subordireciprocates

nate to A (SCHJELDERUPP-EBBE, 1922; ALLEE, 1938). A is the despot and B

afraid of it, adds SCHJELDERUPP-EBBE (1922). According to the
of the way in which dominance
original
descriptions
relationships
is typically

develop (SCHJELDERUPP-EBBE, 1922; ALLEE, 1938), dominance

implies
that individuals
each other.
dominance
status is
recognize
Dyadic
assessed

on the basis of one agonistic interaction

and refers to its consistent, unidirectional
pattern

type, namely
over time.

pecking,

definition":
a slight modification
12. "Modified peck-order
of the "peckorder" definition is to include more than one agonistic interaction
type in
the assessment
of dominance
BARRETTE & VANDAL (1986)
status.
the "peck-order"
definition
of dominance
its
rephrased
summarizing
essence:

dominance

is an attribute

of a relationship
between two individis
uals, whenever an asymmetry in the outcome of agonistic interactions
measured.
If a dominant
seldom has to fight to supplant a subordinate,
the subordinate

is repelled without a need to escalate, then a true domiexists. HAND'S (1986) social dominance
relationship
relationships
between individuals
who meet repeatedly
to this definition.
corresponds
nance

In agonistic dominance
the directionality
of the agonistic
relationships
is not dependent
encounters
upon location, as opposed to territoriality
In
order
to
(KAUFMANN, 1983).
explain this dominance pattern it has been

290
submits at
argued that one member of the dyad in question consistently
the onset of any encounter as a function of the past history of interactions
with the other member
it may be
(BERNSTEIN, 1981). Consequently,
assumed that the subjects discriminate
between different opponents.
A
consistent change in the response pattern of two individuals in agonistic
their first contest,
indicates
that a relationship
conflicts,
following
the
has
been
established.
The
between
defeated member of the
subjects
a
in
submits
or
emits
encounters
dyad
terminating
response
subsequent
with the other member

without

On the basis of such a change

responses in subsequent
agonistic

escalation.

from symmetric to asymmetric

initial
a
with
dominance
can be differentiated
interactions,
dyads
relationship
from those without, since in the latter case each encounter
is contested
before

should not assume that a dominance

yielding. The investigator
exists in each dyad, because unresolved
or egalitarian
relarelationship
&
In
exist
SILK, 1983; HAND, 1986).
(BOYD
tionships may
summary, the
"modified peck-order
definition"
is characterized
by i) asymmetry in the
outcome
of diverse agonistic
avoidance
of escalated
interactions,
ii)
encounters
mediated
and iii)
by the subordinate's
deferring behaviour,

influence

of past encounters
on subsequent
The
responses to opponents.
latter point implies that discrimination
on the basis of individual identity
is a necessary element of dominance
relationships.

Dominance

definition

based

on theoretical

constructs.

The theoretical

to the study of dominance

was introduced
approach
by
HINDE (1978) and GAGE (1981) who postulate
that dominance
must
represent a collection of observable actions, the validity of which can be
between the observables deterempirically verified. The inter-correlation
mines

the

definition

and

usefulness

of the

term

or else justifies

its

rejection.
13. "Intervening
variable":
variable between
intervening

is postulated
in theory as an
variables and a set of depenindependent
dent variables which are intercorrelated
and covary equally in the majorfor examity of dyads (HINDE, 1978; HINDE & DATTA, 1981). Experience,
via the intervening
ple, influences the direction of agonistic interactions
variable

"dominance".

dominance

A dominance/subordinance

relationship

implies

291
one in which

aggression
in the various

is reduced

or (overtly)

absent.

The

interactions,
including non-agonistic
HINDE (1978) uses the term
to the situation.

asymmetry

pattern of
ones, may

vary according
"agressive
as a subcategory
of dominance,
to which other, non-agonisdominance"
behavtic, interactions
may be related. If aggression and non-agonistic
then both can be considered to be under the influence
iour are correlated,
The "intervening
variable"
definition
is also used by
of dominance.
BAENNINGER (1981), CANDLAND & HOER (1981), and GAGE (1981>, among
others.

General

of dominance

attributes

definitions

There

is general agreement
about some potentially
terms
confounding
does not imply
should not be equated to dominance.
Dominance
with
(ALLEE, 1938), and, thus, it is not to be confounded
leadership
in
baboon
"control"
(HINDE, 1978): e.g.
(Papio cynocephalus) consortships,

which

the female may be controlling

as the direction of movement,
female.
The

association

between

between

the different

many aspects of the male's behaviour such

while the male remains dominant
to the
dominance

definitions

and

aggression

of dominance

differs

widely
is
1). Dominance
encounters.
agonistic

(Table

generally associated with conflict resolution during

The "privileged
role" definition of dominance
is an exception, in which
is independent
of any agonistic encounters.
status assignement
Fig. 1
on the way to a definition
shows the minimum set of options encountered
of dominance

which refers to agonistic behaviour.

leads to a different definition.

Each possible

pathway

in the flowchart

are the basis to determine the directionality

When agonistic encounters
of the relationship,
definitions
of dominance
differ with regard to the
In some
expression of overt aggression shown during conflict resolution.
definitions

overt

is the diagnostic
aggression
definitions
do not discriminate

system. Other
and escalated dominance

feature

of a dominance

between non-aggressive
and pool all agonistic encounters to
of
determine status. Yet other definitions explicitly limit the applicability
in
to
those
which
conflict
resolution
the dominance
instances
concept
does not include overt aggression or escalated fights (Table 1 In "peckorder"

definitions,

interactions

overt aggression

by the dominant

individual

may be

292
TABLE 1. Attributes

of common

definitions

of dominance

S = structural definition, F = functional definition, AG = pattern of agonistic interactions

reflects dominance, ASY = asymmetric outcome of contests within a dyad, IR = individual
recognition involved, IH = dominance inheritable, NI =dominance not inheritable, P =
"peaceful" (no overt aggression during conflict resolution), brackets ( ) = not a necessary
condition. LD = low descriptive value, HD = high descriptive value.
the start of an interaction
the default
included

but contests

are generally not escalated, given

These definitions
of the subordinate.
are

yielding response
those in which conflict

among

resolution

is usually a "peaceful"

event.
While

some definitions

consider

dominance

to be an attribute

of the

is aggressiveness"
or "a trait that conveys rank"
("dominance
dominance
as
an
of dyadic relationships
others
view
attribute
definitions),
or of single interactions.
Under definitions which do not regard domi-

individual

nance
dyads

between individuals,
as a relationship
is assigned on the basis of the outcome

subjects
individuals
dominance

are ranked

to the ratio

dominance

status

within

of a single contest or the

of individuals
defeated to

according
lost to. Conversely, under those four definitions which regard
as an attribute
of a relationship
between two individuals,

293

Fig. 1. Flowchart illustrating points of decision when dominance definitions related to

agonistic behaviour are formulated.

294
dominance

status is assigned on the basis of the outcome

of several
if
a
consistent
in
favour
of
one
member
is
conflicts,
asymmetry
dyad
observed
in contest outcome
is a
(Table 1). The dyadic asymmetry
but
not
element
of
the
of
access
to
resources"
possible,
necessary
"priority
definition.
The asymmetry in the direction of
one individual
consistently
winning
consecutive encounters.
Ideally, status
those dyads in which the asymmetry

the outcome
and

of contests

arises from

the other

consistently
losing
should
be
in
made
assignment
only
is statistically significant over a set
is just one possible relationship
relationship

period of time. A dominance

between two individuals.
There are unresolved
and egalitarian
relationin the outcome of contests
ships in which there is no clear asymmetry
A
dominance
between
thus
(HAND, 1986).
individuals,
relationship
meet and resolve conflicts on several
implies that the same individuals
occasions (HINDE & STEVENSON-HINDE, 1976) in a clearly unidirectional
pattern.
The word
definitions.

"relationship"
A dominance

of the individuals

is ambiguous
relationship
concerned.

experience
ALTMANN (1981) and is associated

in the context

of dominance
can be understood
as a mental
This view has been criticized

with those

definitions

by

of dominance

which imply that individual

recognition
operates during interactions
(e.g.
A
use
of the term "relationship"
refers
"peck-order"
definitions).
simpler
between the role of winner and loser
strictly to data, to the interaction
two
individuals
a
series
of agonistic encounters.
The latter
among
during
use corresponds
to the "consistent
winner of agonistic encounters"
and
"the consistent

winner

Some advocates
have

that

at a given context" definitions.

of dominance
as an attribute of dyadic
dominance

relationships

individual

argued
presupposes
recognition
(e.g.
WYNNE-EDWARDS, 1962; ROWELL, 1974; HINDE & STEVENSON-HINDE,
1976). Past conflicts may influence the outcome of future encounters
(e.g.
mice Mus
GINSBURG & ALLEE, 1942; rhesus monkeys Macaca mulatta:
RosE et al., 1972, 1975). Individual recognition,
however, is not a necescondition
to
those
in
cases
which
one
individual consistently
sary
explain
submits to another, familiar individual without an escalated fight taking
place. At the onset of each conflict the opponents may assess one another
on the basis of a morphological
feature or display which correlates well
with competitive
ability (KODRIC-BROWN & BROWN, 1984) regardless of

295
with the same opponent.
This pattern of "peaceful"
past experiences
to the "dominance
is lack of aggressiveconflict resolution corresponds
ness" definition. Individual recognition and memory of past encounters is
cues are not available and
ability when such predictive
fighting ability can only be assessed in escalated contests or inferred from
of the opponent's
behaviour towards other, known members
observation
an important

of the social group.

In conclusion,
individual
produce
definitions,
dominance

is not a necessary condition

to
recognition
observed
under
of
the
dominance
pattern
any
advocates of the "peck-order"
definition envision

the behavioural

hypothesis
of contest

although
as an identity

relationship
use individual

between

two individuals.

that individuals
recognition
success has to be tested in each particular

Dominance

and

The

in their assessment
case.

heritability

can only be inheritable

when it is a property of individuals
studies claim to have selected particularly
(Table 1). Several breeding
status has an inheritable
dominant individuals or to show that dominance
Dominance

component
(e.g. MOORE, 1990; DEWSBURY, 1990; and references therein).
An investigator
can successfully breed for dominant
individuals
on the
basis of the "dominance
is aggressiveness"
definition. Aggressiveness
has
been shown to have an inheritable
in
several
species (e.g.
component
chickens, Gallus domesticus: GUHL et al., 1960; dogs, Canis familiaris: SCOTT
& FULLER, 1965; sticklebacks,
Gasterosteus aculeatus: BAKKER, 1986; silvereyes, Zosterops lateralis: KIKKAWA et al., 1986; fruit flies, Drosophila
melanogaster: HOFFMANN, 1988). On the other hand, breeding experiments
to the determinants
of rank,
suggest that there is a genetic component
when dominance
is defined as an attribute of relationships
(DEWSBURY,
1990).
of the concluand careful formulation
Explicit definition of dominance
sions of breeding experiments
are essential to avoid misunderstandings
about

of
the heritability
of dominance.
Claims regarding the heritability
traits are bound to evoke criticism by the advocates of domi-

dominance

nance as an attribute

of conflict

tion of role. Dominance

ship between

resolution

as an attribute

two individuals

and of dominance
of an interaction

is a relative

measure

as a descripor of a relation-

which

cannot

be

296
selected for (GARTLAN, 1968; ROWELL, 1974; BERNSTEIN, 1981; BARRETTE,
of
properties
1987). Natural selection operates on absolute, competitive
status. Dominance
ranks
and not on relative dominance
individuals
and are, therefore, not an inheritable
on the group composition
and subordiproperty of an individual either (DuNBAR, 1988). Dominants
distinct roles, or
but these are not genetically
nates behave differently,
depend

are fully capastrategies, since subordinates

genetically fixed alternative
circumstances
for a
ble of behaving as dominants
given the appropriate
individuals
in
most
in
status
(MCGUIRE et al., 1984). Furthermore,
change
to some and dominant
to
subordinate
are simultaneously
a hierarchy
of dominance
of the group. The issue of the heritability
of dominance
how a given definition
illustrates
may have applications
with other definitions of the term.
which are incompatible
other

members

and

Descriptive,

predictive

One limitation

of definitions

explanatory

based on observations

that these are not open to empirical investigation.

to determine
proposed, there is no methodology

value

of dominance

rather

than theory
If another definition

is
is

which of the definitions

and the question remains one of semantics rather than

is appropriate
criteria such as the descriptive, prescience (GAGE, 1981). Nevertheless,
between
value can be used to differentiate
dictive and explanatory
definitions.
The

descriptive

value

of dominance.

refers to
value when its definition
has a high descriptive
something for which no other single term existed before and the term is
that something from something
useful when it can be used to differentiate
can be used as a descriptive tool to test the
else. In addition, dominance
Dominance

of behaviour
of merging, and hence simplifying,
categories
of
a dominance
value
(CANDLAND & HOER, 1981). Thus, the descriptive
The
definition is high if it serves efficiently the process of categorization.
of
the
a
of
value
is
an
absolute
definition, irrespective
property
descriptive
to some contentions
context in which it is used. Contrary
theoretical

possibility

value of a
(BERNSTEIN, 1981; HINDE & DATTA, 1981), the descriptive
definition may justify keeping the definition without it being
dominance
Neither the predictive
nor the explanatory
or
explanatory.
predictive

297
value are inherent

qualities of the definition

context in which the concept

absolute

tion of the theoretical

but rather

a func-

is used and of the

concept,
questions asked. In assessing the usefulness of the dominance
one can first estimate its descriptive value, then test empirically whether
the definition is supported by the data, and lastly assess the explanatory
value of using the concept for a particular
question.
In the following definitions,
dominance
has a low descriptive
value
because it is used as a synonym of other, already existing terms, such as
chosen status or role: "domiwinner, or an arbitrarily
aggressiveness,
is dominant,
nance is aggressiveness",
"winner
loser is subordinate",
"successful
definition.

combatant",
status", and "the privileged role"
"reproductive
The use of dominance following these definitions does not help

to describe, predict or explain any phenomena

better than do its synonyms. SYME (1974) argued, with respect to context specific asymmetries
of agonistic interactions,
that if each group has a large
number of "dominance
orders" then the concept is effectively useless. An
in
the
outcome
of particular
interactions
is not a sufficient
asymmetry

in the outcome

a dominance
to introduce
justification
concept, either as a descriptive
mechanism
tool or an explanatory
(DREWS, 1973; SYME, 1974).
The "dominance
is a trait that conveys rank" definition
is used to
characteristics
of an individual
collectively refer to all those behavioural
in
in
are
decisive
the
outcome
of
an
conflict.
Dominance
which
agonistic
value. Dominance
definitions
which
this sense has a high descriptive
include

in the direction
of the outcome of dyadic contests
consistency
and/or lack of aggressive escalation during conflict resolution, also have a
value in that they describe
high descriptive
formerly
unrecognized

phenomena.
In general,

the functional

definition

of dominance,
i.e. "priority
of
access to resources",
describes the access to resources mediated by agonistic contests. Popp & DEVORE (1979) refer more specifically
to the

This definition
equates
pattern of context specific access to resources.
dominance
with non-egalitarian
and consequently
access to resources,
has a high descriptive value since there was no other term to dennote that
phenomenon.
Dominance

in theory language is a hypothetical

construct to investiof certain categories of behaviour (CANDgate the common denominator
LAND & HOER, 1981), and is similar to constructs
such as "stress",

298
"motivation"

and "drives"

(MAXIM, 1981 Dominance

simplifying our description:

cept because it may enable

nance as an intervening
variable between independent
variables, we would reduce the number of links needed
interaction

between

both

sets of variables

is a useful conby using domiand dependent

to describe

the

(HINDE, 1978). Economy in

(HINDE, 1978). When agonistic

is a first step in explanation

description
rank has a high predictive
value in determining
the outcome of other
it is useful to abstract dominance
as a description
of
dyadic relationships,
some aspects of social structure (DEAG, 1977; HINDE, 1978). The descripas an intervening
tive value of dominance
variable is a function of the
between different interaction
types and can be tested
In
female
for
well
inter-correlated
variinstance,
primates,
empirically.
ables could be "supplants",
and "submissive",
"grooming",
possibly with
a weaker link to "priority of access" (HINDE & DATTA, 1981). We know,

inter-correlation

however, that there is lack of consistency even within agonistic categories

since context is likely to influence the pattern observed (Popp & DEVoRE,
variables are unlikely to
1979; LEE, 1983; HAND, 1986). If the dependent
be well correlated
GAGE
the
variable
then, following
(1981),
intervening
can be rejected. The usefulness of dominance,
however, does
between dominance
order and other aspects
not depend on a correlation
of social behaviour.
Dominance
is still useful in the description
of a

definition

particular
type of social structure, in which by some criterion of bossing
related to aggression, the individuals
can be arranged in a hierarchical
order (HINDE,

The

predictive

1978).

value

of dominance.

Dominance

has a high predictive value when the direction or outcome of

can be accurately
a given interaction
on the basis of past
predicted
observations
of dominance
behaviour. Dominance
is, however, a variable
which remains
needs neither to be
dynamic and, thus, predictability
that physical abilities
perfect nor permanent;
ontogeny alone guarantees
"Prediction"
vary (BERNSTEIN, 1981).
(sensu BERNSTEIN, 1981) alludes to
the degree of confidence with which the statistical asymmetry in dyadic
contests

conflict within the

predicts the outcome of any given subsequent
same dyad. Cardinal dominance
ranks have been calculated on the basis
of the probability
of winning contests (BOYD & SILK, 1983). Hence,

299
dominance

is useful as a shorthand

term to indicate

that the outcome of

an agonistic interaction
between two individuals is predictable
with some
level
of
can also be used
practical
certainty (ROWELL, 1974). "Prediction"
to refer to the reliability with which the statistical asymmetry in dyadic
contests

predicts the direction

the context of dominance.

of other,

non-agonistic

interactions

outside

The

value of a dominance
definition
is only of interest,
predictive
when the investigator
uses dominance
to anticitherefore,
relationships
the
behaviour
of
individuals.
The
level
of
of
the
pate
accuracy
prediction
is not an attribute of the definition itself but of the behaviour of the study
animals, and the usefulness of such a prediction depends on the questions
asked in each particular
case. Accurate prediction
alone can be a heuristic value of a dominance
definition without it being explanatory.
This
has yielded contrasting
results, for some authors claim that
approach
dominance
does predict a wide variety of social interactions
(e.g. RICHARDS, 1974; SEYFARTH, 1976, 1980; CHENEY, 1978) and others point out
the lack of correlation
between different measures of dominance
(GARTLAN, 1968; SYME, 1974; BERNSTEIN, 1981; BRAIN, 1981). Variation in the
definitions of dominance
contributes
to the discrepancy
in such results.

The

explanatory

value

It has been contended

behaviour
resources

of dominance.

that the concept of dominance

it should actually predict

that

requires
at least sometimes

as an explanation
priority of access

of
to

(RICHARDS, 1974; DEAG, 1978). This requirea

influences the pattern of
ment, though, implies
priori that dominance
access to resources or else that priority of access to resources be part of
the definition
of dominance.
Predictions
about patterns
of resource
still to be
access, however, are not necessary for the concept of dominance
Dominance
has
a
value
when
the
introducexplanatory.
high explanatory
tion of the term and measurements
of dominance
reduce the magnitude
of formerly unexplained
in the data. The explanatory
variance
value
arises from finding which variables depend on dominance
since, in this
dominance
our
of
social
way,
improves
understanding
organization
to define which is an
(BERNSTEIN, 1981). It is difficult, nevertheless,
measure of dominance
and which are its dependent
independent
ables (BERNSTEIN, 1981). The explanatory
value can be an inherent

variqual-

300
definitions of dominance
but is not
ity of theoretical
(see next paragraph)
a necessary property of structural definitions. It is rather a function of the
theoretical
context in which the concept is used and of the questions
asked by the investigator.
With regard to theoretical

definitions
of dominance,
HINDE (1978)
the
of
dominance
as
an
variable has
argued
concept
intervening
value if the following conditions are satisfied: (1) the investiexplanatory
is
concerned
with more than one dyad, (2) each dyad has a
gation
interactions
multiplex relationship
involving comparable
complementary
that

in which

one bosses the other, and (3) the pattern of inbalance

in those
interactions
is similar in the different dyads. The usefulness of dominance
increases with the variety of correlations
variables
among dependent
(HINDE & DATTA, 1981). In contrast, CANDLAND & HOER (1981) argued
as an intervening
variable is a descriptive
tool which
serves the process of categorization
and is not explanatory.
Dominance
as
an intervening
variable does not make any inferences about the direction
of causality in the observed
correlations
variables.
among dependent

that

dominance

Moreover,

dominance
variable.

as an intervening
variable is, by definition, not an
it
cannot
be incorporated
into an
Consequently,

empirical
and
empirical model to attempt to improve the fit between predictions
dominance
an
variable
can
be
a
observations.
as
Thus,
intervening
powerful descriptive
tool but is of low explanatory
value with respect to
questions about the nature of the observed behaviour.

The

(in)adequacy

of dominance

definitions

Much

of the controversy
around the concept of dominance
has arisen
from the variety of definitions and usages given to the term. Most of the
definitions of dominance
are structural and as such, they are all justified
on semantic grounds (GAGE, 1981). Parsimony,
however, dictates that
there should be an upper practical limit to the number
created for the same term and that their justification
their descriptive
reduce

of new definitions

depend on
to be more useful and to

power. Ideally, for the concept

all users of dominance
should
ambiguity,

should

refer

to one and the

same phenomenon.
In the following discussion I evaluate the adequacy of
the dominance
definitions
listed above and synthesize the essence of a
studies. The criteria used to
unitary dominance
concept for behavioural

301
and less appropriate
into appropriate
are their
respective descriptive values and their affinities to the original definition
as described by SCHJELDERUPP-EBBE (1922). The original
of dominance
categorize

the definitions

definition is not only taken as a reference because it happens to be the

first definition, but also because it has a high descriptive value and called
It is noteworthy that
our attention to a distinct behavioural
phenomenon.
SADE (1981)
in view of the controversy
around dominance
definitions,
which is clear
suggested to look back at the early literature on dominance,
in the definitions and use of the concept (e.g. ALLEE,
and straightforward
1938; COLLIAS, 1944).
listed above, including
the original
Most definitions
of dominance
have in common that they refer to agonistic
definition,
"peck-order"
Dominance
should be reserved to its already widely estabbehaviour.
in which the direction of compleuse for patterns of interaction,
in which
and/or
is
established
through
aggression,
mentarity
initially
Its
usefulness
is
potentially
present (HINDE, 1978).
aggressive behaviour
in interactions,
should not be diluted with other cases of complementarity

lished

e.g. as referred to by WILSON (1975) with respect to food passage in honey

bees. For this reason, as well as because of their low descriptive value, the
status" definitions of dominance
role" and the "reproductive
"privileged
as a
which refer to dominance
are inadequate.
Also, those definitions
or "successful combatant"
are
of aggressiveness,
winner/loser
should
on the basis of their low descriptive value. Dominance
inadequate
not be defined as a synonym of any other already existing term.
BAENNINGER (1981) understands
dominance
as a trait which combines
synonym

several

different

intercorrelated

(see HINDE, 1978), and explicof individuals.

Similarly, WILSON'S

variables

as a property
itly defines dominance
mention
of
dominance
as equivalent to aggressiveness
implies that
(1975)
Definitions
of
dominance
as a
is a variable, individual trait.
dominance
trait of individuals
which
regard

conveys
dominance

"dominance
is a trait
is aggressiveness",
("dominance
with
those
definitions
which
rank") are incompatible
as an attribute

of dyadic

encounters

or relationships,

definition.

including SCHJELDERUPP-EBBE'S "peck-order"

Discontent with the "dominance
is aggressiveness"
by several

emphasized
refers to the patterning
of aggressive
amount

definition

has been

It is generally agreed that dominance

of interactions
between individuals and not to the
authors.

behaviour

that

they

show

(e.g. JAY,

1965;

302
CHALMERS & ROWELL, 1971; ROWELL, 1974;

SYME,
1974; BERNSTEIN, 1976; 1981; HINDE, 1978). Rather, aggression can be
viewed as an expression of dominance
(BERNSTEIN, 1976). Whilst domiit does not
nance may depend on aggression or potential aggressiveness,
correlate
with
them
(LocKWOOD, 1979; BERNSTEIN, 1981). In a
necessarily
KAUFMANN, 1967;

the absolute level

principal factor analysis of wolf (Canis lupus) behaviour,
factor detected
of agonistic behaviour
was not part of the dominance
and communication
(LocKWOOD, 1979). The results of a dominance
analysis in children support
emphasizes the relationship
behaviour

of some group

that
conception of dominance,
coercive
individuals rather than the

the theoretical
between
members

(CAMRAS, 1984).
is known to vary between individuals
(e.g.
aggressiveness
Although
GUHL et al 1960), the levels of aggression observed in natural agonistic
contests do not necessarilly reflect these differences. The game theoretical
to the study of aggressive
(e.g. PARKER, 1974;
competition
approach
&
CARYL, 1980; MAYNARD-SMITH, 1982; ENQUIST LEIMAR, 1990) suggests
that all individuals
observed

have the potential

is a circumstantial

to express

aggression, but that the

of the cost/benefit
ratio

product
aggression
used to
of aggression
of using it. Hence, the directionality
commonly
an
status
is
not
related
to
dominance
aggressive propnecessarilly
assign
but rather to the fact that some individerty of the individuals concerned,
uals defer upon being attacked
contest are low. A definition
several

conflicts

relates

definition

than

dyadic

"peck-order"
definition.
In the previous

because

their chances

of dominance
more
the

based

of winning a given
on the outcome of

to SCHJELDERUPP-EBBE'S (1922)
"dominance
is aggressiveness"

were conthose definitions of dominance

paragraphs,
at all,
which were not related to agonistic interactions
sidered inadequate
as a
had a low descriptive
value because they referred to dominance
as an
synonym of an already existing term, and/or described dominance
inherent

attribute

of individuals

rather

than

of interactions

between

as an attribute
of
to dominance
in
mind
are
now
discussed,
bearing
patterns (Table 1)
that SCHJELDERUPP-EBBE's definition is characterized
by the asymmetry in
and the fact that conflicts are generally
the outcome of dyadic encounters
resolved by the deference of the loser without escalation. The only defini-

them.

conflict

Those

definitions

which

refer

resolution

tions which

regard

asymmetry

and lack of escalation

as necessary

ele-

303
ments

of dominance

are the original "peck-order"

definition
and the
"modified peck-order
definition".
Definitions which consider one of these
conditions as necessary include the "dominance
is lack of aggressiveness"
"consistent
winner of agonistic
contests"
(lack of escalation),
(asymand
the
"consistent
winner
at
contexts"
definition (asymmetry),
given
metry). Asymmetry and lack of escalation may or may not be given in the
conflicts

considered

in the "priority
more noticeably

of access

to resources"

definition,
from the "peck-order"
definition.
which, thus, departs
The "priority of access to resources" definition of dominance
has been
subject to criticism (e.g. SYME, 1974; BERNSTEIN, 1981; BARRETTE & VANon advantages
to dominants
leads to frame a
DAL, 1986). Attention
definition of dominance
based on priority of access to incentives,
while
in favour of consequences
ignoring structure
(BERNSTEIN, 1981). Such
functional
definitions
are problematic
when different
strategies
yield
similar benefits (examples in BERNSTEIN, 1981). If dominance
is measured
on access to only some, specific incentives it is then of no explanatory
value to understand
the principles of social organization.
The use of the "priority of access to resources"
definition is of limited
value to the study of dominance,
because dominance
more generally
refers to all conflict resolution instances, while not all conflicts are over
resources (HAND, 1986). Further, studies using the "priority of access to
resources"

definition

of dominance

of dominance

sometimes

claim to describe

an effect

on resource

dominance

access, a circular statement which arises from

on the basis of the pattern of resource access (e.g.

measuring
GAUTHREAUX, 1978; SYME, 1974; APPLEBY, 1980). This functional definition is, hence, inadequate
to study whether high dominance
rank per se is
The
of an aggressive
order must be
advantageous.
interpretation
restricted
to aggressive behaviour
unless it can be correlated
with a
demonstration

of general priority
of access to resources
when measuring
dominance
on the basis of
(SYME, 1974). In addition,
access to incentives, there is sometimes the problem of identifiying
what
are incentives
of the "priority
of
(BERNSTEIN, 1981). The constraints
access to resources"
definition
illustrated
do
not
above, nevertheless,
make it less valid with respect to the "peck-order"
definition. The "priorquantitative

is concerned
with a
ity of access to resources"
definition,
however,
kind of interaction,
particular
namely contests over resources, irrespective of a dyadic consistency
in their outcome
and of the degree of

304
involved.

escalation

the dominance

respect

For this reason, this definition is incompatible

with
A
similar
of
ScHJELDERUPP-EsBE (1922).
concept

variable"
which also
definition,
argument
applies to the "intervening
lacks affinity with the original definition of dominance.
HINDE &
definition.
Few authors have objected to the "peck-order"
is assessed in terms
DATTA (1981) argued that if dominance/subordinance
variable (here: pecking), then dominance is merely
of only one dependent
would be merely a
and
has
no
value. Dominance
explanatory
descriptive
I
the
data
in
other
words.
argued above that the
way of describing
value is not a property of the definition but of the context in
explanatory
which it is used. Clearly, the original definition was concerned with only
one kind of interaction,
namely pecking, which would limit its applidefinition"
to
other
taxa.
The
"modified
includes
peck-order
cability
interactions
in an analogous
way to
agonistic
treatment
This
modified
definition
also
SCHJELDERUPP-EBBE'S
of pecking.
as a consisderives its high descriptive value from referring to dominance
other

complementary

tent pattern in dyadic interactions

whereby deference rather than escawhich deserves a description
on
lated fights are the rule, a phenomenon
in
are
the
its own. Memory and individual
recognition
implicit
"peckorder"

as proximate
mechanisms
to explain the deference
in
cases but need not be
mechanisms
some
may apply
for the consistent deference behaviour to be observable. Mech-

definition

behaviour.

These

necessary
anisms and function
instead

within

should

hypotheses

not form
concerning

and used
part of the definition
the causation
of dominance

relationships.

The

feature

recognition

criterion

and

dominance

relationships

do not preclude the study of dominance

sexes
and
sizes,
age categories, and even between groups,
or species (e.g. GAUTHREAUX, 1978). SEYFARTH (1981) argued,
populations
that
it makes little functional sense to lump together all age-sex
however,
classes in studying dominance,
because we know about the variety of
Broad

definitions

of dominance

across different

selective presures acting on each class. If the outcome of contests

or a relational attribute which
can be predicted by location (territoriality)
different

correlates

with the direction

useful to describe

of outcomes (e.g. size, age, sex) then it is not

each of them as a special type of agonistic dominance

305
a view tacitly implies that dominance
relation(BERNSTEIN, 1981). Such
does not identify one single reliable
ships exist as long as the investigator
From
of
contest
outcome.
this
dominance
relationperspective,
predictor
ignorance about the identity and interrelaships reflect the investigator's
tion of determinants
of conflict outcomes and dominance
would cease to
exist when these are understood.
This example emphasizes
the imporof explicitly specifying exactly how dominance
is defined (or perif
dominance
is viewed as the
each
However,
ceived) by
investigator.
consistent asymmetry in the outcome of contests between two individuals
tance

whereby the subordinate

generally defers to the dominant without escalaof this pattern is a necessary step
tion, then identifying the determinants
towards

explaining

it and no identified

determinant

should invalidate

the

phenomenon.
In a rigorous

of the "peck-order"
definition of dominance,
application
and individual
determine
the
memory of past encounters
recognition
in dominance
a
behaviour
Such
definition
a
dyads.
implies
relationship
between each of two individuals
and thus, it refers to dominance
as a

in
than just asymmetrical
outcomes
specific phenomenon
non-escalated
contests within a dyad. When observed agonistic patterns
are equivalent to those expected by a peck-order
we need to
dominancc,
know whether identity of the opponent is used to predict the outcome of a
dominance
is inferred.
This
contest, before a peck-order
relationship
much

more

the first contests between

question can be approached
by witnessing
unfamiliar
individuals
and then observing the corresponding
change in
In
over
time.
stable
social
agonistic response
groups, changes in status
instead of
suggest that identity recognition
may be used by opponents
predictive

morphological
as dominance

frequently
ALTMANN (1981)

features

when

such features

do not change

as

status.

regards dominance
relationships as cognitive experiences which should be reserved to the higher primates and humans. He
argues that as long as it is not shown that individuals react to a relationship, rather than to agonistic behaviour
past or present or correlations
need not be inferred. If a
dictates that relationships
parsimony
general feature of the opponent
(like sex or age) is used at the onset of
to decide whether the contest should be escalated
each encounter
or
thereof,

avoided

then past experience

and its identity would be
of the opponent
irrelevant. The feature-recognition
criterion is probably used during most

306
dominant/submissive
once, as explained

interactions
between individuals
which meet only
in the "dominance
is lack of aggression"
definition.

BERNSTEIN (1981) regards the dominant

individual as taking an active
when it acknowledges
immedirelationship
part in the dyadic dominance
ate submission
rather than attacking. There is no need to postulate an
on that ground, if the dominant
individual follows
identity relationship
the simple rule of not attacking when the opponent submits immediately
(a behavioural
feature). Popp & DEVORE (1979) discuss the adaptive value
of stopping further attack once the opponent has submitted, in contexts in
which a relationship
between the two individuals is irrelevant.

Individuals,

sex,

groups,

or species:

the

units

of dominance

The concept of dominance

has been mostly utilized to characterize
the
It has been also applied to the relationship
behaviour
of individuals.
such as sexes, groups, and species. The working
between
categories
in studies
of intersexual
dominance
(e.g. RALLS, 1976;
hypothesis
DESROCHERS Bt al., 1988) is that one sex may exclude the other from
resemlimiting resources. Such an exclusion is referred to as dominance,
of access to resources"
functional
definition.
Domibling the "priority
nance is not to be interpreted
as the agent causing resource monopolizais measured on the basis of the outcome of all
tion because dominance
most of which may be over resources. Dominance
agonistic encounters,
in this context is descriptive,
not explanatory.
in
dominance
is similar to the dominance
Intergroup
relationships
dyads
entire

of familiar

because
individuals,
behaves as a unit which

in intergroup
contests often the
is known to the opponent
group

group
dominance
in wild rhesus monkeys
from previous encounters.
Intergroup
on
the
of
has been described
basis
supplants, avoidance, or fights between
at
groups (SOUTHWICK et al., 1965). This usage of the term corresponds,
the

group
definition.

level,

to

the

"consistent

winner

of

agonistic

contests"

dominance
after MORSE (1974) is the priority of access to
Interspecific
resources that results from successful attacks, fights, chases or supplanting
actions present or past. It represents
one mechanism
of interference
For example, gulls (Larus sp.) dominate
curlews (Numenius
competition.
while
curlews
dominate
small
et
waders
arquata),
(ENS al., 1990). WAGNER

307
& GAUTHREAUX(1990) refer to MoRSE's (1974) definition as the definition
in general. GAUTHREAUX (1978) discussed the ecological
dominance following the "priority of access to
significance of interspecific

of dominance

can be
definition. Other claims that the concept of dominance
to
and
have
referred
species (WILSON, 1975)
applied
groups, populations
loser is subordinate",
or the "dominance
is
to the "winner is dominant,
Such definitions were shown above to be
lack of aggression"
definitions.
resources"

and the same reasoning

at the level of individuals,
inadequate
to units of higher order.
their application
A further
outside

applies

to

of applying the concept of dominance

to interactions
social group is that this use is not compatible
with
of the term that imply a relationship
between the interactants.
of the latter definition argue that dominance
hierarchies
are

problem
the familiar

definitions
Advocates

is a prereqonly possible in small groups because individual aquaintance

uisite (WYNNE-EDWARDS, 1962; ROWELL, 1974). In the usage of dominance beyond the confines of the familiar group, the individual identity of
the opponent and past agonistic experiences with that particular
individIn interspecific
ual are often irrelevant to the decisions of the contestants.
dominance

interactions
the dyadic conflict is probably settled by recogniof
attributes
tion
of the opponent.
species specific
The crucial criterion to justify the use of dominance
for higher order
the
choice
of
rather
definition
is
not
but
the
of
categories
category,
dominance

used.

interactions

between

of dominance

is used beyond the level of dyadic

single conspecifics there is the danger that analogies
will be confused with homology (BERNSTEIN, 1981).).

The

of dominance:

essence

When

dominance

synthesis

and

definition

it is generally agreed
definitions,
Despite a great variety of dominance
that dominance
refers to agonistic behaviour.
Several definitions of dominance were rejected on the grounds of their low descriptive
value. The
original definition of dominance
by SaHJEt,nERUPP-EssE (1922) described
a distinct phenomenon
in chicken, which is regarded here as the basis for
a more general concept of dominance
with broader
The
applicability.
high descriptive value of this dominance
concept justifies its use; prediction and explanation
are applications
of dominance,
which originate in
The
field of interest of each investigator.
essence of domithe particular

308
nance

emerges from the above discussion

structural
definition of the term:

which

following,

is summarized

in the

is an attribute

of the pattern of repeated,

agonistic interactwo individuals,
characterized
a
consistent
outcome in
by
favour of the same dyad member and a default yielding response of its
rather than escalation.
The status of the consistent winner is
opponent
and that of the loser subordinate.
dominant
Dominance

tions between

definitions of dominance,
based on the above definition, can
Operational
in each particular
be formulated
study according to the choice of subjects
and the theoretical
framework
used. Although
this definition
concenit
trates on dyadic interactions
(because these are readily observable),
does not preclude

the possiblity that the outcome is influenced

by the
nearby presence of other individuals, which may tacitly or overtly support
to the
one of the contestants
(see KAWAI ( 1958, 1965) for an introduction
rank).
concept of dependent
A semantic clarification
this definition: dominance
status
complements
and dominance
ranks are different measures (HINDE, 1978; BERNSTEIN,
BARRETTE
& VANDAL, 1986; DUNBAR, 1988). This distinction
1984;
which refer to the pattern
of
Dominance
status
refers
to
a
dyadic
(Table 1).
relationship
between two animals and depends on an animal's attributes compared to
those of the other, while rank refers to the relative position within a group
to the definitions

applies

of dominance

encounters

Dominance
status refers to
and, hence, depends on group composition.
the status of one individual
within a given dyad and can be either
dominant

or subordinate

to the direction
of the statistically
according
in the outcome of several contests. Dominance

significant
asymmetry
rank refers to the position of one individual in a dominance
hierarchy and
in greek letters (sex
can be expressed at an ordinal level either numerically,
=
=
Q
bottom
rank, SCHJELDERUPP-EBBE, 1922) or qualitatively
top rank,
as high or low, but not as dominant
or subordinate.
In general, dominance
ranks are calculated after assessment of dominance

status

in all possible dyads of the group. The resulting hierarchy

can be linear, but does not necessarilly have to be so (APPLEBY, 1983).
hierand the dominance
Dyadic dominance/subordinance
relationships
archy are not necessarily

related

and the first does not imply the existence

309
of ordinal
in a social group (HINDE, 1978). The limitations
distance
between
ranks
is the
the
definition
ranks,
adjacent
by
to
indices
which
reflect
the
same, have lead some investigators
develop
distances"
that separate animals in a hiermore realistic "dominance
ARCESE
& LUDWIG, 1986; ZUMPE &
BOYD
&
SILK, 1983;
archy (e.g.
of the other

in which

MICHAEL, 1986; DREWS, in prep.).

has been viewed from
With respect to the above definition, dominance
is seen as a descripAt a general level dominance
different perspectives.
tive tool (e,g. CANDLAND & HOER, 1981). ALTMANN (1981) and SEYFARTH
and dominance
hierarchies
relationships
(1981) maintain that dominance
have no function

and are not ends in themselves

or cognitive

experiences
structural descriptive terms used by
of the animals, but simply shorthand,
can be regarded
the observer. BERNSTEIN (1981) argued that dominance
as a relative measure used to illustrate one attribute of a dyadic relationas a useful estimate of an
relationships
ship. SMUTS(1981) sees dominance
ability to influence
through that individual's

individual's

the behaviour

of another

for its own

benefit,
power to inflict physical injury. Popp &
as an unavoidable
DEVORE (1977) view agonistic dominance
product of
cost/benefit
considerations
They, conseduring competitive interactions.
under
the
of
for
frame
dominance
aggressive
theory
games
quently,
competition.
Other investigators

concentrate

on the fact that

fights are
of the communi-

escalated

as a dimension
generally avoided and regard dominance
cation system (e.g. CHANCE, 1956; MAXIM, 1981). The subordinate
may
completes the attack or immediately
signal defeat before the dominant
when intention to attack is signaled by stereotyped movements or vocalizthat a communication
system has evolved
in
the
that
The
lack
of
way
investigators
perceive
agreement
(SADE, 1981).
or the other
for one interpretation
is evident, and preference
dominance
is bound to remain arbitrary.
The structural definition proposed above simplifies the debate around
ations.

This is seen as evidence

about the menconcept because it makes no assumptions

of
us
who use
about
those
of the subjects or
tal experiences
investigators
the concept. This definition provides a frame for future studies on dominance which can deal with more specific aspects, such as the role of
the dominance

memory
minants

as well as the identification

and individual recognition,
of dominance
status.
and consequences

of deter-

310
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