Академический Документы
Профессиональный Документы
Культура Документы
Brill, Leiden
ANIMAL BEHAVIOUR
by
CARLOS DREWS1)
(Department of Zoology, Cambridge University, Downing Street, Cambridge CB2 3EJ,
U.K.)
(With 1 Figure)
(Ace. 7-VII-1993)
Abstract
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 neces
sarilly 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 domi
nance 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
1) I thank S.K. ELTRI~<GHAM, G.W. NORTON, and P.C. LEE for valuable discussion.
284 CARLOS DREWS
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 individ
ual (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
tion neither implies that escalation is avoided nor that individuals recog
nize each other and incorporate past agonistic experiences into contest
decisions.
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,
THE CONCEPT AND DEFINITION OF DOMINANCE 293
DEFINITION AS
THEORETICAL CONSTRUCT
F1JNCTIONAL STRUCTURAL
I
V
v v
v
CONTEXTS EACH CONTEXT TREATED
SEPARATELY
OBSERVED PATTERN
OF INTERACTIONS
CALLED DOMINANCE
RELATIONSHIP
DOMINANCE DEFINITION
value are inherent absolute qualities of the definition but rather a func
tion of the theoretical context in which the concept is used and of the
questions asked. In assessing the usefulness of the dominance concept,
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
aggressiveness, winner, or an arbitrarily chosen status or role: "domi
nance is aggressiveness", "winner is dominant, loser is subordinate",
"successful combatant", "reproductive status", and "the privileged role"
definition. The use of dominance following these definitions does not help
to describe, predict or explain any phenomena better than do its syn
onyms. SYME (1974) argued, with respect to context specific asymmetries
in the outcome of agonistic interactions, that if each group has a large
number of "dominance orders" then the concept is effectively useless. An
asymmetry in the outcome of particular interactions is not a sufficient
justification to introduce a dominance concept, either as a descriptive
tool or an explanatory mechanism (DREWS, 1973; SYME, 1974).
The "dominance is a trait that conveys rank" definition is used to
collectively refer to all those behavioural characteristics of an individual
which are decisive in the outcome of an agonistic conflict. Dominance in
this sense has a high descriptive value. Dominance definitions which
include consistency in the direction of the outcome of dyadic contests
and/ or lack of aggressive escalation during conflict resolution, also have a
high descriptive value in that they describe formerly unrecognized
phenomena.
In general, the functional definition of dominance, i.e. "priority of
access to resources", describes the access to resources mediated by ago
nistic contests. Por-e & DEVORE (1979) refer more specifically to the
pattern of context specific access to resources. This definition equates
dominance with non-egalitarian access to resources, and consequently
has a high descriptive value since there was no other term to dennote that
phenomenon.
Dominance in theory language is a hypothetical construct to investi
gate the common denominator of certain categories of behaviour (CAND
LAND & HOER, 1981), and is similar to constructs such as "stress",
298 CARLOS DREWS
categorize the definitions into appropriate and less appropriate are their
respective descriptive values and their affinities to the original definition
of dominance as described by SCH1ELDERUPp-EBBE (1922). The original
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
our attention to a distinct behavioural phenomenon. It is noteworthy that
in view of the controversy around dominance definitions, SADE (1981)
suggested to look back at the early literature on dominance, which is clear
and straightforward in the definitions and use of the concept (e.g. ALLEE,
1938; COLLIAS, 1944).
Most definitions of dominance listed above, including the original
"peck-order" definition, have in common that they refer to agonistic
behaviour. Dominance should be reserved to its already widely estab
lished use for patterns of interaction, in which the direction of comple
mentarity is initially established through aggression, andf or in which
aggressive behaviour is potentially present (HINDE, 1978). Its usefulness
should not be diluted with other cases of complementarity in interactions,
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
"privileged role" and the "reproductive status" definitions of dominance
are inadequate. Also, those definitions which refer to dominance as a
synonym of aggressiveness, winnerfloser or "successful combatant" are
inadequate on the basis oftheir low descriptive value. Dominance should
not be defined as a synonym of any other already existing term.
BAENNINGER (1981) understands dominance as a trait which combines
several different intercorrelated variables (see HINDE, 1978), and explic
itly defines dominance as a property of individuals. Similarly, WILSON'S
(1975) mention of dominance as equivalent to aggressiveness implies that
dominance is a variable, individual trait. Definitions of dominance as a
trait of individuals ("dominance is aggressiveness", "dominance is a trait
which conveys rank") are incompatible with those definitions which
regard dominance as an attribute of dyadic encounters or relationships,
including SCH1ELDERUPp-EBBE'S "peck-order" definition.
Discontent with the "dominance is aggressiveness" definition has been
emphasized by several authors. It is generally agreed that dominance
refers to the patterning of interactions between individuals and not to the
amount of aggressive behaviour that they show (e.g. JAY, 1965;
302 CARLOS DREWS
Literature cited
ALLEE, W.C. (1938). The social life of animals. - W.vV.Norton, New York.
ALTMANN, S.A. (1981). Dominance relationships: the Chcshire cat's grin? - Behav. Brain
Sci. 4, p. 430-431.
ApPLEBY, M.C. (1980). Social rank and food access in red dcer stags. - Behaviour 74, p.
294-309.
- - (1983). The probability of linearity in hierarchies, - Anim.Behav. 31, p. 600-608.
ARCESE, P. & LUDWIG, D. (1986). Improving estimates of dominance based on ratios. -
Condor 88, p.106-107.
BAENNINGER, R. (1981). Dominance: on distinguishing the baby from the bathwat er. -
Bchav. Brain Sci. 4, p. 431-432.
BAKKER, T.C.M. (1986). Aggressiveness in sticklebacks (Gasterosteus aculeatus L.): a bchav
iour-genetic study. -- Behaviour 98, p. 1-144.
BARRETTE, C. (1987). Dominance cannot be inhcrited. - TREE 2: 251
- - & VANDAL, D. (1986). Social rank, dominance, antler size, and access to food in snow
bound wild woodland caribou. - Behaviour 97, p. 118-146.
BERNSTEIN, LS. (1976). Dominance, aggression and reproduction in primate societies.
J. theor.Biol. 60, p. 459-472
- - (1981). Dominance: The baby and the bathwater. - Behav. Brain Sci. 4. p. 419-457.
- - (1984). The adaptive value of maladaptive behavior, or you've got to be stupid in
order to be smart. - Ethol.Sociobiol. 5, p. 297-303.
BOYD R. & SILK, J.B. (1983). A method for assigning cardinal dominance ranks.
Anim.Behav. 31, p. 45-58.
BRAIN, P.F. (1981). The concept of dominance also has problems in studies on rodents.
Behav. Brain Sci. 4, p. 434-435.
CA~lRAS, L.A. (1984). Children's verbal and nonverbal communication in a conflict situa
tion. - Ethol.Sociobiol. 5, p. 257-268.
CANDLAND, K.D. & HOER, J.B. (1981). The logical status of dominance. - Behav. Brain
Sci. 4, p. 436-437.
CARYL, P.G. (1980). Escalated fighting and the war of nerves: gamcs theory and animal
combat. - In: Perspectives in ethology 4-. (P.P.G. BATESON & P.H. KLOPFER, eds).
Plenum Press, New York, p. 199-224.
CHALMERS, N.R. & ROWELL, T.E. (1971). Behavior and female reproductive cycles in a
captive group of mangabeys. -- Folia primatol. 14, p. 1-14.
CHANCE, M.R.A. (1956). Social structure of a colony of Macaca mulatta.
BritJ.Anim.Behav. 4, p. 1-13.
CHENEY, D.L. (1978). Interactions of immature male and female baboons with adult
females. - Anim.Behav. 26, p. 389-408.
CLUTTON-BROCK, T.H. & HARVEY, P.H. (1976). Evolutionary rules and primate societies.
- In: Growing points in ethology (P.P.G. Bateson & R.A. HINDE, eds). The University
Press, Cambridge, p.195-237.
- - , ALBON, S.D., GIBSON, R.M. & GUINESS, F.E. (1979). The logical stag: adaptive
aspects of fighting in red deer (Cervus elaphus L.). - Anim.Behav. 27, p. 211-225.
COLLIAS, N.E. (1944). Aggressive behavior among vertebrate animals. - Physiol. Zool. 17,
p.83-123.
DEAG,J.M. (1977). Aggression and submission in monkey societies. - Anim.Behav. 25, p.
465-474.
- - (1978). The adaptive significance of baboon and macaque social behaviour. - In:
Population control by social behaviour. (P. EBLINGAAND & D.M. STODDART, eds).
Institute of Biology. London, p. 83-ll3.
DESROCHERS, A., HANNON, SJ. & NORDIN, K.E. (1988). Winter survival and tcrritory
acquisition in a northern population of black-capped chickadees. - The Auk 105, p.
727-736.
THE CONCEPT AND DEFINITION OF DOMINANCE 311
DEWSBURY, D.A. (1982). Dominance rank, copulatory behavior, and differential reproduc
tion. - QRev.Biol. 57, p. 135-159.
- - (1990). Fathers and sons: genetic factors and social dominance in deer mice, Per
omiscus maniculatus. - Anim.Behav. 39, p. 284-289.
DREWS, D.R. (1973). Group formation in captive Galago crassicaudatus: notes on the domi
nance concept.- Z. Tierpsychol. 32, p. 425-435.
DUNBAR, R.UvL (1988). Primate social systems. - Croom Helm, London & Sydney.
EBERHARD, MJ.W. (1969) The social biology of polis tine wasps. - Miscellaneous Publica
tions, Museum of Zoology, University of Michigan, Ann Arbor 140, p. 1-10 I.
ENQUIST, M. & LEIMAR, O. (1990). The evolution of fatal fighting. - Anim.Behav. 39, p.
1-9.
ENS, B., ESSELINK, P. & ZWARTS, L. (1990). Kleptoparasitism as a problem of prey choice: a
study on mudflat-feeding curlews, Numenius arquata. -- Anim.Behav. 39, p. 219-230.
FEDIGAN, L. (1982). Primate paradigms: Sex roles and social bonds. - Eden Press,
Montreal.
GAGE, F.H. (1981). Dominance: measure first and then define. - Behav. Brain Sci. 4, p.
440-441.
GARTLAN, j.S. (1968). Structure and function in primate society. - Folia primatol. 8, p.
89-120.
GAUTHREAUX, S.A.jr. (1978). The ecological significance of behavioural dominance. - In:
Perspectives in ethology 3. (P.P.G. BATESON & P.H. KLOPFER, eds). Plenum Press, New
York, p. 17-54.
GINSBURG, V. & ALLEE, W.C. (1942). Some effects of conditioning on social dominance and
subordination in inbred strains of mice. -- Physiol. Zool. 15, p. 485-506.
GUHL, A.M., CRAIG,j.V. & MUELLER, C.D. (1960). Selective breeding for aggressiveness in
chickens. - Poultry Sci. 39, p. 970-980.
HANBY, j. (1976). Sociosexual development in primates. - In: Perspectives in ethology 2.
(P.P.G. BATESON & P.H. KLOPFER, eds). Plenum Press, New York, p. 1-67.
HAND, J.L. (1986). Resolution of social conflicts: dominance, egalitarianism, spheres of
dominance, and game theory. - QRev.Biol. 61, p. 201-220.
HINDE, R.A. (1978). Dominance and role - two concepts with dual meaning.
j.soc.bioI.Struct. I, p. 27-38.
- - & DATTA, S. (1981). Dominance: an intervening variable. - Behav. Brain Sci. 4, p.
442.
- - & STEVENSON-HINDE,J. (1976). Towards understanding relationships: dynamic stabil
ity. - In: Growing points in ethology. (P.P.G. BATESON & R.A. HINDE, eds). Cam
bridge University Press, New York, p. 451-479.
HOFFMANN, A.A. (1988). Heritable variation for territorial success in two Drosophila
melanogaster males. - Anim.Behav. 36, p. 1180-1189.
JAY, P. (1965). The common langur of North India. -- In: Primate behavior, field studies of
monkeys and apes. (1. DEVORE, ed.). Holt, Rinehart and Winston, New York, p.
197-249.
KAWAI, M. (1958). On the system of social ranks in a natural group ofJapanese monkeys,
(parts I & II). - Primates I, p. 111-148 (in Japanese).
- - (1965). On the system of social ranks in a natural group ofJapanese monkeys: Basic
rank and dependent rank. - In: japanese monkeys: A collection of translations (K.
IMANISHI & S.A. ALTMANN, eds). Emory University Press, Atlanta, p. 66-86.
KAUFMANN, J.H. (1967). Social relations of adult males in a free-ranging band of rhesus
monkeys. - In: Social communication among primates. (S.A. ALTMANN, ed.). Univer
sity of Chicago Press, Chicago and London: p. 73-98.
- - (1983). On the definitions and functions of dominance and territoriality. - Biol.Rev.
58, p. 1-20.
312 CARLOS DREWS
KIKKAWA, L., SMITH,J.N.M., PRYS-JONES, R., FISK, P. & CATTERALL, C. (1980). Determi
nants of social dominance and inheritance of agonistic behavior in an island popula
tion of silvereyes (;:;:,osterops lateralisi. - Behav.Eco!.Sociobio!. 19, p. 165-169.
KODRIC-BROWN, A. & BROWN,J.H. (1984). Truth in advertising: the kinds of traits favoured
by sexual selection. - Am. Nat. 124, p. 309-323.
VAN KREVELD, D. (1970). A selective view of dominance- subordination relations in mam
mals. - Genetic Psycho!. Manogr. 81, p. 141-173.
LEBoEUF, B.]. (1974). Male-male competition and reproductive success in elephant seals.
- Am. Zoo!' 14, p. 163-176.
LEE, P.C. (1983). Context specific unpredictability in dominance interactions. - In:
Primate social relationships. (R.A. HINDE, cd.). Blackwell Scientific Publications,
Oxford, p. 35-44.
LOCKWOOD, R. (1979). Dominance in wolves: useful construct or bad habit? - In: The
behaviour and ecology of wolves. (E. KLINGHAMMER, ed.). Garland Press, New York, p.
225-244.
MAXIM, P.E. (1981). Dominance: a useful dimension of social communication. - Behav.
Brain Sci. 4, p. 444-445.
MAYNARD-SMITH, J. (1982). Evolution and the theory of games. - Cambridge University
Press, Cambridge.
MCGUIRE, M.T, RALEIGH, M.J. & BRAMMER, G.L. (1984). Adaptation, selection, and
benefit-cost balances: implications of behavioral-physiological studies of social domi
nance in male vervet monkeys. - Etho!.Sociobio!. 5, p. 269-277.
MOORE, A.J. (1990). The inheritance of social dominance, mating behaviour and attrac
tiveness to mates in male Nauphoeta cinerea. - Anim.Behav. 39, p. 388-397.
MORSE, D.H. (1974). Niche breadth as a function of social dominance. - Am. Nat. 108, p.
818-830.
PARKER, G.F. (1974). Assessment strategy and the evolution of fighting behaviour.
j. theor.Bio!. 47, p. 223-243.
Por-r-, J.L. & DEVORE, 1. (1979). Aggressive competition and social dominance theory:
synopsis. - In: The great apes. (D.A. HAMBURG & E.R. MCCOWN, eds). The Ben
jamin/Cummings Publishing Co., Menlo Park, p. 317 -338.
RALLS, R. (1976). Mammals in which females arc larger than males. - QRev.Bio!. 51, p.
245-276.
RICHARDS, S.M. (1974). The concept of dominance and methods of assessment.
Anim.Behav. 22, p. 914-930.
ROSE, R.M., GORDON, TP. & BERNSTEIN, 1.S. (1972). Plasma testosterone levels in the male
rhesus: influences of sexual and social stimuli. - Science, 178, p. 643-645.
- - , BERNSTEIN, LS. & GORDON, T.P. (1975). Consequences of social conflict on plasma
testosterone levels in rhesus monkeys. - Psychosom. Medicine, 37, p. 50-61.
ROWELL, TE. (1974). The concept of social dominance. - Behav. Bio!. II, p. 131-154.
SADE, D.S. (1981). Patterning of aggression. - Behav.Brain Sci. 4,p. 446-447.
SCHJELDERUPp-EBBE, T. (1922). Be itrage zur Sozialpsychologie des Haushuhns.
Zeitsch.f.Psycho!. 88, p. 226-252.
SCOTT, J.P. & FULLER, J.L. (1965). Genetics and the social behavior of the dog.
University of Chicago Press, Chicago.
SEYFARTH, R.M. (1976). Social relationships among adult female baboons.
Anim.Behav., 24, 917-938.
- - (1980). The distribution of grooming and related behaviors among adult female
vervet monkeys. - Anim.Behav. 28, p. 798-813.
- - (1981). Do monkeys rank each other? - Behav. Brain Sci. 4, p. 447-448.
SMUTS, B. (1981). Dominance: an alternative view. ~- Behav. Brain Sci. 4, p. 448-449.
SOUTHWICK, C.H., BEG, M.A. & SIDDIQI, M.R. (1965). Rhesus monkeys in North India.
In: Primate behavior. Field studies of monkeys and apes. (L DEVORE, cd.). Holt,
Rinehart and Winston, New York, p. 111-159.
THE CONCEPT AND DEFINITION OF DOMINANCE 313