Review of the concept and definition of dominance in animal behaviour.
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The concept of dominance has contributed greatly to our understanding of social structurc 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 thc usefulness and mcaning 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) emergcd as the basis to formulate a structural definition with wide applicability and which reflects the essencc of the concept: Dominance is an attribute of the pattern of rcpcated, 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 dominancc, 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 greatly to our understanding of social struc
ture, Tables listing dominance studies in a diversity of animal taxa are
1) I thank S.K. ELTRI~<GHAM, G.W. NORTON, and P.C. LEE for valuable discussion.
284 CARLOS DREWS
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 regarding its meaning. SCH]ELDERUPP-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 of domestic fowl, the dynamics of the hierarchy, and the
determinants of rank. \VILSON (1975) summarized different types of domi
nance hierarchies found in animals. HArm (1986) provided an ample
discussion of the different categories found among dominance relation
ships and rigorously defined the terms commonly used in studies of
dominance. Other conceptual discussions about dominance can be found
in VAN KREVELD (1970), FEDIGAN (1982) and DUNBAR (1988).
A lively debate has underlined the application of dominance in behav
ioural studies over the past three decades (e.g. BERNSTEIN, 1981). GARTLAN
(1968) criticized the different interpretations of dominance because these
often led to ambiguity. He further criticized the fact that the term
dominance is often used without a definition or else is arbitrarily
redefined to fit the findings. ROWELL (1974) pointed out that the defini
tion of dominance is elusive. Investigators are far from a consensus on
how to define and measure dominance (BERNSTEIN, 1981), leading to
disparate opinions regarding its function and evolution. BERNSTEIN (1981)
concentrated on primates but his review of the problems associated with
the concept of dominance has general application. His article is accom
panied by comments from various investigators and thus provides an
overview of the dominance debate and lack of concensus at that time.
One of the central problems of the dominance debate has been that
critics have not always explicitly specified which definitions they advocate
and which they criticize. This has kept the debate going since the criti
cisms may have been justified with respect to a particular definition but
not to others. In this paper, an overview of the diversi ty of definitions of
dominance found in the animal behaviour literature is presented and the
usefulness of each definition assessed with respect to furthering our
understanding of social structure and behaviour. Elimination of some
definitions and finding the common denominator of others lead to a
synthesis illustrating the essence of dominance in behavioural studies.
This synthesis is condensed in the definition of dominance proposed in
the concluding section.
285 THE CONCEPT AND DEFINITION OF DOMINANCE
Categories of d.ornfnance definitions
The terms winner, loser, submissive, subordination and aggression are
part of the specialized vocabulary surrounding the dominance concept.
These terms are defined and used here as follows:
Winner: the contestant that expresses consummatory behaviour according to its initial goal (BAENNINGER, 1981; HAND, 1986). Winner/loser refers to the outcome of one single contest.
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).
Dominance definitions can be separated into those based on empirical
observations and definitions based on theoretical constructs (GAGE, 1981,
HINDE & DATTA, 1981). These have been referred to as "data-language"
and "theory-language" definitions respectively by HINDE & DATTA
(1981). GAGE (1981) distinguishes two approaches to the study of domi
nance: in the first, the investigator formulates in theory-language the
conditions which must be met by a useful application of the term domi
nance, then derives a testable hypothesis and lets the results of empirical
inquiry support the definition of dominance or else lead to its rejection. In the second approach, the investigator assumes the existence of domi
nance, asserts an operational definition, and proceeds to demonstrate the
utility of the term and definition. This latter approach is similar to the
data-language definitions mentioned by HINDE & DATTA (1981), in which
a particular behavioural interaction is given the name "dominance inter
action". More broadly, dominance is used to describe a particular kind or
set of observable interactions. The dominance definitions based on
empirical observations can in turn be subdivided into structural and
that individuals recognize each other. Dyadic dominance status is
assessed on the basis of one agonistic interaction type, namely pecking,
and refers to its consistent, unidirectional pattern over time.
12. "Modified peck-order definition": a slight modification of the "peck
order" definition is to include more than one agonistic interaction type in
the assessment of dominance status. BARRETTE & VANDAL (1986)
rephrased the "peck-order" definition of dominance summarizing its
essence: dominance is an attribute of a relationship between two individ
uals, whenever an asymmetry in the outcome of agonistic interactions is
measured. If a dominant seldom has to fight to supplant a subordinate,
the subordinate is repelled without a need to escalate, then a true domi
nance relationship exists. HAND'S (1986) social dominance relationships
between individuals who meet repeatedly corresponds to this definition.
In agonistic dominance relationships the directionality of the agonistic
encounters is not dependent upon location, as opposed to territoriality
(KAUFMANN, 1983). In order to explain this dominance pattern it has been
290 CARLOS DREWS
argued that one member of the dyad in question consistently submits at
the onset of any encounter as a function of the past history of interactions
with the other member (BERNSTEIN, 1981). Consequently, it may be
assumed that the subjects discriminate between different opponents. A consistent change in the response pattern of two individuals in agonistic
conflicts, following their first contest, indicates that a relationship
between the subjects has been established. The defeated member of the
dyad submits or emits a terminating response in subsequent encounters
with the other member without escalation. On the basis of such a change
from symmetric to asymmetric initial responses in subsequent agonistic
interactions, dyads with a dominance relationship can be differentiated
from those without, since in the latter case each encounter is contested
before yielding. The investigator should not assume that a dominance
relationship exists in each dyad, because unresolved or egalitarian rela
tionships may exist (BOYD & SILK, 1983; HAND, 1986). In summary, the
"modified peck-order definition" is characterized by i) asymmetry in the
outcome of diverse agonistic interactions, ii) avoidance of escalated
encounters mediated by the subordinate's deferring behaviour, and iii)
influence of past encounters on subsequent responses to opponents. The
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 approach to the study of dominance was introduced by
HINDE (1978) and GAGE (1981) who postulate that dominance must
represent a collection of observable actions, the validity of which can be
empirically verified. The inter-correlation between the observables deter
mines the definition and usefulness of the term or else justifies its
rejection.
13. "Intervening variable": dominance is postulated in theory as an
intervening variable between independent variables and a set of depen
dent variables which are intercorrelated and covary equally in the major
ity of dyads (HINDE, 1978; HINDE & DATTA, 1981). Experience, for exam
ple, influences the direction of agonistic interactions via the intervening
variable "dominance". A dominance/ subordinance relationship implies
THE CONCEPT AND DEFINITION OF DOMINANCE 291
one in which aggression is reduced or (overtly) absent. The pattern of
asymmetry in the various interactions, including non-agonistic ones, may
vary according to the situation. HINDE (1978) uses the term "agressive
dominance" as a subcategory of dominance, to which other, non-agonis
tic, interactions may be related. If aggression and non-agonistic behav
iour are correlated, then both can be considered to be under the influence
of dominance. The "intervening variable" definition is also used by
BAENNINGER (1981), CANDLAND & HOER (1981), and GAGE (1981), among
others.
General attributes of dOininance definitions
There is general agreement about some potentially confounding terms
which should not be equated to dominance. Dominance does not imply
leadership (ALLEE, 1938), and, thus, it is not to be confounded with
"control" (HINDE, 1978): e.g. in baboon (Papio cynocephalusi consortships,
the female may be controlling many aspects of the male's behaviour such
as the direction of movement, while the male remams dominant to the
female.
The association between dominance and aggression differs widely
between the different definitions of dominance (Table 1). Dominance is
generally associated with conflict resolution during agonistic encounters.
The "privileged role" definition of dominance is an exception, in which
status assignement is independent of any agonistic encounters. Fig. 1
shows the minimum set of options encountered on the way to a definition
of dominance which refers to agonistic behaviour. Each possible pathway
in the flowchart leads to a different definition.
'When agonistic encounters are the basis to determine the directionality
of the relationship, definitions of dominance differ with regard to the
expression of overt aggression shown during conflict resolution. In some
definitions overt aggression is the diagnostic feature of a dominance
system. Other definitions do not discriminate between non-aggressive
and escalated dominance interactions and pool all agonistic encounters to
determine status. Yet other definitions explicitly limit the applicability of
the dominance concept to those instances in which conflict resolution
does not include overt aggression or escalated fights (Table 1). In "peck
order" definitions, overt aggression by the dominant individual may be
292 CARLOS DREWS
TABLE 1. Attributes of common definitions of dominance
Ernpdr-ica! observation definitions
Non-agonistic behaviour: Dominance as a role description: - "privileged role"
Agonistic behaviour as part of definition: - "reproductive status" Dominance as a property of the individual: - "dominance is aggressiveness" - "dominance is a trait that conveys rank" Dominance as an attribute of dyadic encounters: - "winner is dominant, loser is subordinate" - "successful combatant" - "priority of access to resources" - "dominance is lack of aggressiveness"
Consistent outcome of dyadic interactions: - "consistent winner of agonistic contests" - "consistent winner at a given context"
Escalation usually avoided: - original "peck-order" - "peck-order" modified
S, AG, LD, IH NT S, AG, LD S, AG, LD F, (AG), (ASY), HD S, AG, P, HD S, AG, ASY, HD
S, AG, ASY, HD S, AG, ASY, IR, P, HD S, AG, ASY, IR, P, HD S, AG, ASY, P, HD
AG, NI, HD/LD
S = structural definition, F = functional definition, AG = pattern of agonistic interactions reflects dominance, ASY = asymmetric outcome of contests within a dyad, TR = 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 but contests are generally not escalated, given
the default yielding response of the subordinate. These definitions are
included among those in which conflict resolution is usually a "peaceful"
event.
While some definitions consider dominance to be an attribute of the
individual ("dominance is aggressiveness" or "a trait that conveys rank"
definitions), others view dominance as an attribute of dyadic relationships
or of single interactions. Under definitions which do not regard domi
nance as a relationship between individuals, dominance status within
dyads is assigned on the basis of the outcome of a single contest or the
subjects are ranked according to the ratio of individuals defeated to
individuals lost to. Conversely, under those four definitions which regard
dominance as an attribute of a relationship between two individuals,
293 THE CONCEPT AND DEFINITION OF DOMINANCE
IBASIS: AGONISTIC BEHAVIOUR
~ ~
DEFINITION AS THEORETICAL CONSTRUCT
F1JNCTIONAL STRUCTURAL
I V
ESCALATED CONTESTS INCLUDED OR EXCLUDED FROM DEFINITION
v v
OUTCOME OF ONE CONTEST CONSISTENT OUTCOME FROM SEVERAL IS SUFFICIENT CONTESTS BETWEEN SAME INDIVIDUALS
CONTEXTS
RELATIONSHIP
v
EACH CONTEXT TREATED SEPARATELY
OBSERVED PATTERN OF INTERACTIONS CALLED DOMINANCE
DOMINANCE DEFINITION
Fig. I. Flowchart illustrating points of decision when dominance definitions related to agonistic behaviour are formulated.
294 CARLOS DREWS
dominance status is assigned on the basis of the outcome of several
conflicts, if a consistent asymmetry in favour of one dyad member is
observed (Table I). The dyadic asymmetry in contest outcome is a
possible, but not necessary element of the "priority of access to resources"
definition.
The asymmetry in the direction of the outcome of contests arises from
one individual consistently winning and the other consistently losing
consecutive encounters. Ideally, status assignment should only be made in
those dyads in which the asymmetry is statistically significant over a set
period of time. A dominance relationship is just one possible relationship
between two individuals. There are unresolved and egalitarian relation
ships in which there is no clear asymmetry in the outcome of contests
(HAND, 1986). A dominance relationship between individuals, thus
implies that the same individuals meet and resolve conflicts on several
occasions (HINDE & STEVENSON-HINDE, 1976) in a clearly unidirectional
pattern.
The word "relationship" is ambiguous in the context of dominance
definitions. A dominance relationship can be understood as a mental
experience of the individuals concerned. This view has been criticized by
ALTMANN (1981) and is associated with those definitions of dominance
which imply that individual recognition operates during interactions (e.g. "peck-order" definitions). A simpler use of the term "relationship" refers
strictly to data, to the interaction between the role of winner and loser
among two individuals during a series of agonistic encounters. The latter
use corresponds to the "consistent winner of agonistic encounters" and
"the consistent winner at a given context" definitions.
Some advocates of dominance as an attribute of dyadic relationships
have argued that dominance presupposes individual 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 sp.: GINSBURG & ALLEE, 1942; rhesus monkeys Maraca mulatta:
ROSE et al., 1972, 1975). Individual recognition, however, is not a neces
sary condition to explain those cases in which one individual consistently
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 THE CONCEPT AND DEFINITION OF DOMINANCE
past experiences with the same opponent. This pattern of "peaceful"
conflict resolution corresponds to the "dominance is lack of aggressive
ness" definition. Individual recognition and memory of past encounters is
an important ability when such predictive cues are not available and
fighting ability can only be assessed in escalated contests or inferred from
observation of the opponent's behaviour towards other, known members
of the social group.
In conclusion, individual recognition is not a necessary condition to
produce the behavioural pattern observed under any of the dominance
definitions, although advocates of the "peck-order" definition envision
dominance as an identity relationship between two individuals. The
hypothesis that individuals use individual recognition in their assessment
of contest success has to be tested in each particular case.
Domdrrarrce and heritability
Dominance can only be inheritable when it is a property of individuals
(Table 1). Several breeding studies claim to have selected particularly
dominant individuals or to show that dominance status has an inheritable
component (e.g. Nlo0RE, 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 component in several species (e.g. chickens, Gallus domesticus: GUHL et al., 1960; dogs, Canisfamiliaris: SCOTT
(1981) maintain that dominance relationships and dominance hierarchies
have no function and are not ends in themselves or cognitive experiences
of the animals, but simply shorthand, structural descriptive terms used by
the observer. BERNSTEIN (1981) argued that dominance can be regarded
as a relative measure used to illustrate one attribute of a dyadic relation
ship. SMUTS (1981) sees dominance relationships as a useful estimate of an
individual's ability to influence the behaviour of another for its own
benefit, through that individual's power to inflict physical injury. Por-t- &
DEVORE (1977) view agonistic dominance as an unavoidable product of
cost/benefit considerations during competitive interactions. They, conse
quently, frame dominance under the theory of games for aggressive
competition.
Other investigators concentrate on the fact that escalated fights are
generally avoided and regard dominance as a dimension ofthe communi
cation system (e.g. CHANCE, 1956; MAXIM, 1981). The subordinate may
signal defeat before the dominant completes the attack or immediately
when intention to attack is signaled by stereotyped movements or vocaliz
ations. This is seen as evidence that a communication system has evolved
(SADE, 1981). The lack of agreement in the way that investigators perceive
dominance is evident, and preference for one interpretation or the other
is bound to remain arbitrary.
The structural definition proposed above simplifies the debate around
the dominance concept because it makes no assumptions about the men
tal experiences of the subjects or about those of us investigators who use
the concept. This definition provides a frame for future studies on domi
nance which can deal with more specific aspects, such as the role of
memory and individual recognition, as well as the identification of deter
minants and consequences of dominance status.
310 CARLOS DREWS
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