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Review
Dominance in domestic dogs revisited:
Useful habit and useful construct?
Matthijs B.H. Schilder
a
,
*
, Claudia M. Vinke
a
, Joanne A.M. van der Borg
b
a
Department of Animals in Science & Society, Faculty of Veterinary Medicine, Utrecht University, The Netherlands
b
Department of Animal Sciences, Behavioural Ecology Group, Wageningen University, Wageningen, The Netherlands
article info
Article history:
Received 10 December 2013
Received in revised form
8 April 2014
Accepted 8 April 2014
Available online 16 April 2014
Keywords:
dominance hierarchy
domestic dog
formal dominance
humanedog relationship
abstract
In the last decade, the validity and relevance of the dominance model was regularly put into question
regarding relationships between canids like dogs and wolves, and consequently, humanedog
relationships as well. The concept underlying this model, scientically dened as an intervening
variable reecting status difference between individuals, is applicable when formal status signals
symbolize the long-term relationship between individuals, resulting in a formalized dominance hi-
erarchy. This article reviews the basics underlying the concept of dominance and reects on the value
and importance of some new quantitative studies on the applicability of the concept of dominance in
domestic dogs. The conclusions are, rst, that formal dominance is present in the domestic dog,
expressed by context-independent unidirectional formal status signals. Consequently, formal
dominance (e.g., submission) plays an important role in assessing status in dogedog relationships.
Second, that nonverbal statuserelated communication in humans resembles that in dogs to a
considerable degree, and hence dogs may be well able to interpret this human statuserelated
nonverbal communication from their perspective. Dominance is therefore also likely to play a role in
humanedog relationships. Hence, the dominance concept might be useful to explain the develop-
ment of certain problems in dogedog and dogehuman relationships. However, enforcing a dominant
status by a human may entail considerable risks and should therefore be avoided.
Ó2014 Elsevier Inc. All rights reserved.
Introduction
The last decade saw a discussion on the validity and relevance of
the dominance model regarding relationships between canids like
dogs and wolves, and consequently, also concerning the humane
dog relationship. The reasons for this were summarized in a much
discussed article by Bradshaw et al. (2009), which denounces the
concept of dominance in dogs and wolves on several grounds, to be
mentioned in Dominance in dogs: considering the pro and contra
arguments in more detail section. Recently, however, 3 new inde-
pendent quantitative studies conrm the concept of dominance to
be applicable in domestic dogs (Cafazzo et al., 2010; Trisko, 2011;
van der Borg et al., 2012). Moreover, these studies also mention
and conrm the existence of the so-called formal dominance in
dogs. This aspect of dominance was ignored in previous discussions
on dogs, whereas this is well known in primates as a most impor-
tant expression of submission or dominance. One exception is the
study of Bauer and Smuts (2007), who recognize formal dominance
in dogs, partly on the basis of a quantitative analysis of play
behavior. These 3 recent quantitative studies used the research
model developed at Utrecht University in the years 1970-1980 (de
Waal, 1977; van Hooff and Wensing, 1987). Therefore, it is time to
reconsider the arguments, data, and methods leading to such
opposing conclusions and also to clarify whether the recent con-
rmations of dominance in domestic dogs could be a result of a bias
in the methodology.
In this article, we rst establish the theoretical backgrounds for
dominance and its counterpart submission. In doing so, we follow
the logic of the Utrecht School of former professor Jan van Hooff and
his former pupil Frans de Waal, who have signicantly contributed
to the development of the concept of dominance and to the
methodology to investigate what role dominance might play as an
organizing principle in species, and to what degree. Subsequently,
*Address for reprint requests and correspondence: Matthijs B. H. Schilder, PhD,
Department of Animals in Science & Society, Faculty of Veterinary Medicine, Utrecht
University, P.O. Box 80166, 3508 TD Utrecht, The Netherlands. Tel: þ31-30-2533674;
Fax: þ31-30-2539227.
E-mail address: m.b.h.schilder1@uu.nl (M.B.H. Schilder).
Contents lists available at ScienceDirect
Journal of Veterinary Behavior
journal homepage: www.journalvetbehavior.com
1558-7878/$ esee front matter Ó2014 Elsevier Inc. All rights reserved.
http://dx.doi.org/10.1016/j.jveb.2014.04.005
Journal of Veterinary Behavior 9 (2014) 184e191
we will discuss the validity and power of the argumentations and
data from authors opposing the idea of dominance playing a role in
dogs and wolves and contrast these with the results from the recent
studies using the Utrecht dominance model. Finally, we will discuss
possible implications for the humanedog relationship. We do not
discuss in depth the mechanisms of formation of dominance re-
lationships and the connection between dominance and leadership.
The former has been modeled in the studies by Hemelrijk (2002),
Beacham (2003), Forkman and Haskell (2004), and Dugatkin and
Dugatkin (2007), and the latter is discussed in the studies by
Peterson et al. (2002), Bonanni et al. (2010), and Akos et al. (2014).
Some basics of dominance
Members of a social group may differ in many aspects. Differ-
ences may include asymmetries regarding physical power, stamina,
lineage, personality, weight, weaponry, age, and so on (Maynard
Smith and Parker, 1976; Chase and Seitz, 2011). These differences
in personal properties of individuals may inuence the dominance
relationships (Bernstein, 1981) and may be stable over some time
and to a certain degree. Stable asymmetries between individuals
may lead to more or less predictable differences in behavioral
outputs and outcomes of conicts over resources. This predict-
ability is not 100% because other factors (e.g., motivation) may
interfere.
Dominance is regarded as an intervening variable that summa-
rizes a set of behavioral differences between individuals (cf, Hinde,
1974; Hinde and Datta, 1981). This rst means that a number of
relevant behavioral exchanges within each dyad should show
identical main directions: animal A shows several relevant behav-
iors more frequently toward animal B than vice versa. This rst
aspect can be investigated by correlation analysis and subsequent
cluster or principal component analysis. This analysis may reveal
groups of behavioral elements that are performed with the same
main directions within the dyads. Behaviors in such a cluster may
be lumped for further analysis. Typically, when main directions of
submissive behaviors are reversed, these should correlate well with
the behavioral measures reecting dominance.
Second, these main directions within dyads should be identical
over different competitive contexts. That is to say that the main
directions of behaviors within each pair of individuals in context A
should be identical to those in contexts B and C. For example, the
main directions of behavioral exchanges in mate competition should
be identical to those in food competition. An example of this type of
consistency in dogs is given by the study of Cafazzo et al. (2010).
Third, these behavioral asymmetries should be stable over a
considerable period to be measured over weeks, months, and even
several years.
Fourth, if a certain relevant behavior is found to be exchanged
exclusively in one direction (animal A performs behavior X toward
animal B, but [almost] never vice versa), it is said that such a
behavior functions as a formal dominance signal or even a meta-
communicative signal (cf, van Hooff and Wensing, 1987), conveying
either dominance or submission. Such formal dominance signals
have been described in several species including wolves and several
primates (see, e.g., van Hooff and Wensing (1987) for wolves and de
Waal (1977) for Java monkeys). Formal dominance signals should be
shown exclusively in the same direction, independent of the type of
context. There are formal signals that express dominance and
formal signals that express subordination (Preuschoft, 1999). In the
latter case, these signals may also function to inhibit further
aggressive or dominant behavior by dominants. For example, the
silent bared teeth display in Java and Barbary macaques is exclu-
sively shown by submissive individuals toward dominants, inde-
pendent of the context (de Waal and Luttrell, 1985; Preuschoft and
van Schaik, 2000). In wolves, a high posture is shown by dominants
toward subordinates, whereas a low posture is exclusively shown
by subordinates toward higher ranking animals, again independent
of the context (van Hooff and Wensing, 1987).
We will use the term status signals (cf, Flack and de Waal, 2004)
as a shortcut, indicating signals that are involved in either signaling
dominance or submission.
In some articles (e.g., de Waal, 1989), an additional criterion to
recognize formal status signals is added, namely that such a 1-sided
signal should be shown not only during competitive conicts but
also outside conicts, for example, during greeting. This has been
shown to occur in free-ranging domestic dogs also (Bonanni et al.,
2010). The detection of formal status signals in a species is taken
as proof that dominance relationships are important in structuring
a given social organization. However, if formal status signals cannot
be demonstrated, this does not exclude the possibility that domi-
nance does play some role. Rather it suggests that its role might be
far more limited (see, e.g., Schilder, 1988).
The degree of one sidedness by which a certain behavior is
shown can be expressed in an index, called the direction (in)con-
sistency index (van Hooff and Wensing, 1987). For example, a di-
rection consistency index of 90% means that an average of 90% of all
occurrences of that particular behavior in all dyads were shown by
animal A toward animal B and only 10% by animal B toward animal
A. For dogs, formal status signals were suggested by Bauer and
Smuts (2007) in their article on play in dogs. They observed that
role reversalsoccurred during play and involved chasing and
tackles but never included mounts, muzzle bites, and muzzle licks,
suggesting that these behaviors were invariant indicators of formal
dominance during play in domestic dogs (italics ours).
For a human observer, the usefulness of a formal status signal to
identify the rank of individuals may sometimes be hampered by the
rarity of its performance; preferably, it should be shown in most
dyads. This aspect is called the coverage. It may occur that a rare
signal of formal dominance, because of its one-sidedness, could in
principle very well be used as a dominance indicator but that its
rarity prevents its use to clarify rank relationships between a large
number of individuals in a group. When it is shown, however, it can
be extremely meaningful. An example is the behavior head-on-
hindquarters in plains zebra stallions. This is very much a 1-sided
behavior, and its main directions were stable over years, but it
occurred in only a few dyads and was therefore not suited to be
used as a criterion to rank all stallions in a herd (Schilder, 1988).
A last characteristic of dominance in a social group is that
stable relationships often lead to a ranking of individuals, which is
completely or nearly linear. However, triangles may exist, meaning
that animal A dominates animal B, that animal B dominates C,
whereas animal C dominates A. The degree of linearity is depen-
dent on the number of blank relationships (no behavioral ex-
changes for that particular behavior in a dyad), the number of ties
(both A and B perform a behavior to one another with the same
frequency), and the number of triangles. Several procedures to
construct rank order and test their linearity and steepness have
been published (de Vries and Appleby, 2000; de Vries et al., 2000;
Gammell et al., 2001; Bayly et al., 2006). Ranking individuals in a
group may be useful to describe behavioral patterns in that group
but may not (necessarily) reect an important variable in social
organization (Bernstein, 1981).
A quantitative analysis as described previously may lead to a
variety of results: nonconcordant distributions of behaviors over
dyads leading to nonidentical rank orders, or behaviors that are
exchanged reciprocally, show an insufcient coverage, or a
nonsignicant degree of linearity. An example of such an analysis
leading to a limited applicability of the dominance concept as
sketched can be found in the study by Schilder (1988) for plains
M.B.H. Schilder et al. / Journal of Veterinary Behavior 9 (2014) 184e191 185
zebra stallions. Therefore, the methodology is not bound to produce
biased results but is precise and sensitive. The fact that 3 recent
independent studies on dominance in dogs using this methodology
found formal dominance to be present in the dog cannot therefore,
with any likelihood, be attributed to a bias inherent to the
methodology.
Finally, the validation of the concept of dominance must be
found in its ultimate biological consequence, namely reproductive
success. Indeed, the relationship between dominance status and
reproductive success has been found in many species (see Ellis,
1995 for a review; African wild dogs: Creel et al., 2007). Most, but
not all these studies, conrm the biological importance of being
high ranking, and therefore they point at the important role of
dominance relationships. Especially in primatology, the study of
dominance, its causes, and its biological implications has been
developed to great depth. Many of these studies took many years,
enabling assessment of (sometimes lifelong) reproductive output in
relation to rank position. One such study has been performed on
free-ranging dogs. The authors found that, although the pack
comprised multiple breeding individuals, both male copulation
success and female reproductive success were positivelyinuenced
by a linear combination of dominance rank, age, and leadership
(Cafazzo, personal communication). Spotte (2012), on the other
hand, summarizes that neither rank nor body size are important
factors inuencing reproductive success in the dog. Doubt is cast
concerning this conclusion as, according to many articles, rank
orders are not based on formal status signals.
It has become clear that dominance relationships become
established not because higher ranking individuals reinforce their
status by being aggressive or showing formal dominance signals
but because lower ranking individuals recognize supremacy by
showing formal submissive signals (e.g., Rowell, 1974; Syme, 1974;
van Hooff and Wensing, 1987). This statement also makes sense
from a logical point of view: if acceptance is lacking, conicts are
bound to remain or individuals avoid each other and may leave
their group (see Mech and Cluff, 2010 for wolves). A state of
repeated conicts may occur when perceived asymmetries be-
tween 2 individuals are too small. In turn, this explains why con-
icts between some dogs in the same household may be more
frequent, persistent, and erce than between other combinations.
The probability of conicts should be increased when dogs of the
same sex, same age, and same breed are involved, leading to min-
imal asymmetry and hence, to more uncertainty with regard to
their relationship. With this point, we now arrive at the assumed
short-term function of dominance, namely to restrain the number
and severity of physical conicts. In canids, we know of just 2
studies that show that a stable hierarchy decreases and an instable
situation increases the occurrence of aggressive encounters in
semicaptive wolves (Zimen, 1975; Moran 1982). Nevertheless, it
may occur that either the submissive animal does not recognize its
status or the dominant animal will still reinforce its status, in spite
of the submissive animal showing submissive behavior (Forkman
and Haskell, 2004; Mech and Cluff, 2010).
In principle, conicts are costly because they consume energy
and time, may lead to wounds and other risks, such as an increased
vulnerability to predation as a consequence of a decreased atten-
tion toward predators. These are the reasons why individuals have
to make an a priori assessment before entering into a conict. This
assessment would include processing information on whether the
resource at stake is worth the risk, the chances of losing and win-
ning, and the possible costs of entering into a conict. The outcome
of such a weighing should lead to a decision to ght or to refrain and
retreat. Uncertainty concerning the resource holding potential
(RHP) could lead to agonistic interactions. These provide in-
dividuals with opportunities to gain information on the strength of
opponents, which consequently can result into avoiding ghts with
animals that could defeat them. In this way, physical harm or worse
may be reduced (de Waal, 1989).
To form dominance relationships resulting in a rank order, ani-
mals do not need to have a concept of dominance, which is sug-
gested by Casey (2009). It sufces that dogs know who is superior
and who is subordinate, and a self-organized rank order emerges
(Beacham, 2003). Moreover, unpublished data from Utrecht Uni-
versity showed that dogs also have insight into the relationships
between third parties. If this were not true, interventions in on-
going interactions would be randomly directed. However, the
data showed that they are clearly consistent with the rank order!
Third-party interventions turned out to be more directed against
lower than higher ranking individuals in an interacting pair and
more against losers than winners (Netto et al., 1992). Also, trying to
obtain a higher position does not need future planning, as Casey
(2009) suggests. It just may happen provided there is some moti-
vational factor that might be inuenced by genetic (family) re-
lationships, health, and environmental factors (see Chase and Seitz,
2011). That many domestic dogs, if not most, do not show such an
ambitionmay be also the result of domestication. In this respect,
individual and breed differences are to be expected. Also, breeds
may differ in their possibilities to show submissive and other be-
haviors (Goodwin et al., 1997), and breed-specic tail and ear
postures may inuence communicative (im)possibilities (Mertens,
2004; Leaver and Reimchen, 2008).
Dominance in dogs: considering the pro and contra
arguments in more detail
In this part, we follow the main arguments that would invalidate
the concept of dominance for domestic dogs and comment on
ndings and conclusions.
1. Personality trait or dimension of relationship? Some authors
claim that dominance is not a trait, but a characteristic of a
relationship (Bernstein, 1981; Langbein and Puppe, 2004;
Bradshaw et al., 2009). Consequently, use of the expression
dominant dogis meaningless(Bradshaw et al., 2009, p. 138)
Comparable statements can be found in psychology. For
example, Burgoon and Hale (1984) dene dominance and
submission as fundamental dimensions of personal relation-
ships, whereas others recognize dominance as a personality trait
(Kalma, 1991; Gangestad et al., 1992; Zebrowitz and Collins,
1997). There has thus been considerable debate in human
and dog personality research on how to score and interpret
certain traits and over the classication of certain behaviors
within trait spectrums (e.g., Jones and Gosling, 2005; DeYoung
et al., 2013). For instance, submissiveness has been classied as
a trait in its own right (Jones and Gosling, 2005) and also as a
subtrait of neuroticism (Ley et al., 2008).
Dominance, like subordination, reveals itself in interactions
between individuals, but it is the differences between personal
characteristics that dene nature and outcomes of these in-
teractions in terms of dominance relationships (see also Bernstein,
1981, p. 422). There is no logical reason why a trait like dominance/
submission would not be present in canids and other social species.
Data from studies on dog personality up to now were not linked to
data on dominance in social groups. An attempt was made by Akos
et al. (2014), who investigated leadership, dominance, and per-
sonality in a small group of dogs. These authors state on page 2 that
leader/dominant dogs have a unique personality: they are more
trainable, controllable and aggressive, additionally they are older
than follower/subordinate dogs.
M.B.H. Schilder et al. / Journal of Veterinary Behavior 9 (2014) 184e191186
Gosling and John (1999) compared 19 studies in 12 species
regarding the 5-factor model of personality. They included domi-
nance and activity as factors in their analysis and concluded that
evidence for a dominance factor was modest. Dominance appeared
as a separate factor in 7 of the 19 studies they reviewed. In animals,
the dominance factor was typically dened by assertiveness and
boldness, physical aggression, and low fearfulness (p. 71). In
humans, the dominance trait is typically only related to the E factor:
extraversion vs. introversion (Gosling and John, 1999). Most likely
dominant dogs will be of the boldtype as dened by Svartberg
(2005). This behavioral tendency often cannot be demonstrated at
an early age in dogs or wolves (dogs: Beaudet et al., 1994; Diederich
and Giffroy, 2006; wolves: Packard, 2003), although in a recent
meta-analysis Fratkin et al. (2013) found substantial consistency for
the dimensions aggression and submission in puppies, just as
Svartberg et al. (2005) found consistency in personality in adult
dogs. Inconclusive as the results are, they suggest that aspects of
dominant and submissive personalities can be recognized in per-
sonality studies, including those on dogs. There are indeed dogs
that show high postures toward many other dogs, just as there are
dogs that show submissive behaviors toward many other dogs. This
may reect respectively dominant and submissive personalities,
which may be recognized by many other individuals, although bold
(certain) and shy (uncertain) would also be appropriate labels.
2. Explaining submissive actions as conict defusing actions, instead
of submissive ones, leaves the one-sidedness of the performance
of submissive behaviors unexplained: why should only sub-
missive dogs defuse and not the dominant ones? There must be
more to it than just reconciliation or appeasement. This does
not mean that acknowledgment of the dominant status of
the opponent is the only function of showing a submissive
behavior. Because muzzle licking seems to be derived from
juvenile begging behavior (Schenkel, 1947), both a submissive
and an appeasing meaning could have been developed later on
in evolution. This defusing function of submissive acts was
recognized by Schenkel (1967). The one-sidedness of submis-
sive behaviors, nevertheless, demonstrates that acknowledg-
ment of status might be a primary function.
3. The much used pack model of wolves does not apply for domestic
dogs; a conclusion that was drawn by Coppinger (2001), van
Kerkhove (2004), Bradshaw et al. (2009), and Yin (2009).
In wild wolves, life appears far more peaceful than previously
thought, and wolves form an extended family group (Mech, 1999;
Mech and Boitani, 2003). Under natural circumstances, strict rank
orders, separately between male and female wolves, were not
found in the wild and ghts about dominance were not observed by
Mech (1999).Mech (1999) seems to equal alleged dominance by the
breeding pair with the parenting role: any parent is dominant to
its young offspring,so the dominance status is trivial informa-
tion,and the term dominance falsely implies a rigid force-based
dominance hierarchy.However, Mech (1999) may have over-
looked here that between the breeding male and female, and also
within the litter, dominance relationships may exist and depend on
formal submissive signaling. On the other hand, Mech (1999) rec-
ognizes that there is rank-related communication in wild wolves
and that it involves posturing. Unfortunately, he has neither
appreciated nor investigated whether posturing could be a formal
status signal in wild wolf packs and that its frequent occurrence
suggests that it must be functional. In a later article, Mech and Cluff
(2010) described a prolonged dominance interaction between 2
wolves. Based on his observations that dominant wolves, usually
the parents in a family group, dominate offspring by forcing them to
the ground (mostly not lasting longer than 30 seconds), he suggests
that adult wolves this way force mature offspring to leave the pack
and disperse. Research by him was done in summer, and com-
petition for food and mates in summer is less than in winter. This
may explain why he has missed much of dominance-related
communication.
Most data on dominance in wolves stem from groups in
captivity, where group composition may be abnormal and where
levels of competition and aggression may be higher because of the
lack of opportunities to disperse as wild wolves have. Consequently,
increased frequency and intensity of communicative acts facilitate
the construction of rank orders by observers. Because competition
arises mostly within sexes, resulting in far more interactions within
rather than between sexes, separate rank orders may arise. Such
hierarchies were not found in all studies on captive wolves: Moran
(1982, p. 81) did not nd an overall social structurein his captive
group of wolves, although there were pairs in the group whose
interactions suggested a typical dominance/subordinate relation-
ship.Spotte (2012, p. 227) concluded that expressions of domi-
nance hierarchies in wolf packs are largely artifacts of captivity.
However, a more adequate view would be that the species has the
property to form hierarchies when competition is increased. There
are no reports of abnormal behaviors involved in this, so formation
of dominance relationships can be seen as a normal process
occurring in extreme conditions resulting in elevated competition.
Further arguments that the wolf model cannot be applied to
dogs are rst, that dogs do not live in wolf-like cooperative packs
and second, that dog behavior differs radically from wolf behavior
because of domestication (Bradshaw et al., 2009). The studies by Pal
et al. (1998) and Pal et al. (1999) are cited by Bradshaw et al. to
illustrate these differences (see later). More recent, however, a clear
pack structure in free-ranging dogs has been revealed (Bonanni
et al., 2010; Cafazzo et al., 2010, 2012). Although not all feral dogs
may adopt a wolf-like pack structure, the data in the newer articles
bear out that communication regarding status does not differ so
much between wolves and dogs at all. The main results of the new
studies on dogs are summarized in Table 1 and resemble closely
Table 1
Signals related to (formal) dominance in wolves, dogs, and humans
Authors Formal signals of dominance and
submission in the wolf
Van Hooff and Wensing, 1987 Low & high posture,
muzzle licking
Fatjo et al., 2007 Tail position
Mech, 1999 Posturing
Schenkel, 1947, 1967 Passive submission
Authors Formal signals of dominance and
submission in the dog
Bauer and Smuts, 2007 Mounts, muzzle licks, muzzle bites
Cafazzo et al., 2010 Submissive afliative behavior
including low posture
Trisko, 2011 Submissive behavior
van der Borg et al., 2012 Lowering of posture, muzzle licks,
low tail wag
¼active submission cf
Schenkel, 1967
Authors Signals of dominance and
submission in humans
Morris, 1979; Mignault & Chaudhuri,
2003
Lowering of posture: crouch, kneel,
head down, kiss hand/foot
Zivin, 1977; Schwartz et al., 1982;
Eibl-Eibesfeldt, 1997; Carney et al.,
2005
High head/body posture
Rosa and Masur, 1979; Kalma, 1989 Extended eye contact
Strongman and Champness, 1968 Gaze aversion
Bold print: signals designated by authors as formal signals; normal print: domi-
nance/submission signals not designated as formal signals by respective authors.
M.B.H. Schilder et al. / Journal of Veterinary Behavior 9 (2014) 184e191 187
those in the classic descriptive study of Schenkel (1947) and the
quantitative study by van Hooff and Wensing (1987) on wolves, and
are also in line with the study by Fatjo et al. (2007), who concluded
that the most reliable indicator of status is the tail position. It
should be noted that the captive group studied by van Hooff and
Wensing (1987) in fact was an extended family: an original
breeding pair with young and adult offspring.
Although the behavior of captive wolves may differ quantita-
tively from that of free-ranging wolves related to the level and
intensity of competition, so far, the available data show that the
principles of communication are identical. Because formal status
signals in wild wolves do occur, dominance must also play a role
there. That feral dogs do not always live in clear pack structures
might complicate discovery of dominance relationships but does
not inherently makes them nonexistent. The argument that a wolf
group must be seen as an extended cooperative family does not
logically imply that dominance does not play an important role:
data on humans and other primates (who often live in an extended
family group structure!) show otherwise (Tiger, 1970; de Waal,
1977; Thomas, 2012). Mech and Cluff (2010) even stated that
dominance is one of the most pervasive and important behaviors
among wolves in a pack, yet its signicance in free-ranging packs
has been little studied.
4. The outcomes of competition about resources have been used to
investigate and dene dominance. It is important to note the
often overlooked difference between dominance and the con-
sequences thereof. For example, dominance has been dened by
van Kerkhove (2004), following Drews (1993):with reference
to repeated conicts between conspecics over a scarce
resource, whereby the same individual always gains access to
the resource. With respect to only that specicresource, the
winning animal is said to be dominant over the loosing animal.
Thus, social dominance is often quite uid .(italics ours). In
this reasoning, dominance is confused with a possible conse-
quence of being dominant, and second, the implication is that
dominance is exible. Used in this way, the concept of domi-
nance indeed would be superuous. We have argued before
that using the concept of dominance only makes sense if
relationships tend to be stable over time and its behavioral
expression consistent over contexts. This also is likely to hold
for canids: Fatjo et al. (2007) mention a stable rank order in
their captive wolf group lasting at least 12 months and com-
parable stability was found in the group of wolves studied by
van Hooff and Wensing (1987) and later by Derix et al. (1993).
Flexibility in the outcomes of conicts may well be explained
by individual differences in motivation and/or perception of the
incentive value of a resource, leading to outcomes inconsistent
with the dominance relationship. For example, a satiated
dominant individual may leave food for a submissive one, but
formal dominance signals will be exchanged as they were
before: their main directions will not change. When formal
dominance is not taken into account, and the focus of domi-
nance studies is on the outcomes of conicts only, this may lead
to false assumptions regarding dominance status. Competitive
ability is considered to differ from formal dominance (de Waal,
1989; Preuschoft, 1999).
5. In feral dogs, reproduction is not linked with dominance. Pal et al.
(1999, 2003, 2005) could neither nd clear displays of domi-
nance or submission in their group of feral dogs nor frequent
aggressive encounters between females. They found little
reproductive suppression in females, and they found that
infanticide only occurred rarely. However, Pal et al. (1998)
established dominance hierarchies based on aggressive en-
counters, which is supposedly not an adequate measure, as
discussed before. Moreover, Pal et al. (1999) could not detect a
relationship between dominance and the number of matings.
From this, Bradshaw et al. (2009) concluded that reproduction
in feral dog groups .appears not to be controlled by a wolf
pack type of dominance hierarchy.It can be questioned if such
a conclusion can be made based on data on this small group of
animals; and it certainly should not be generalized. It remains
also unclear from their article if Pal et al. (1998) have investi-
gated whether asymmetries in behavioral exchanges are
concordant before lumping behaviors into groupings for
investigating dominance and its consequences. In a later study,
Pal (2010) refers to Cafazzo et al. (2010) regarding their dening
criteria for dominance in litters of feral dogs: he now used
upright and stiff body postures with head held high and ears
pricked, and growling and baring teeth.In this study, he states
that dominance hierarchy develops at an early stage of life and
that female and male domestic dogs form intra-sexual domi-
nance relationships(p. 150). This result agrees with those on
wolves of Schenkel (1947), Woolpy (1968), and Zimen (1975)
but not with that of van Hooff and Wensing (1987).
Data analysis in several species has shown that lumping be-
haviors that are not distributed in an identical way may prevent
nding existing rank relationships to emerge. For example,
combining offensive with defensive aggression in horses masks an
otherwise linear rank order (van Dierendonck et al., 1995). Thus,
Palsndings might be simply a matter of a small data set, data
arrangement, and analysis. As mentioned before, a recent study by
Cafazzo et al. (personal communication) has shown a relationship
between dominance and reproductive success in free-ranging dogs.
6. Dogs are not likely to strive for dominance.Bradshaw et al. (2009)
and Casey (2009) presented some data of their own research,
on a group of neutered male dogs. In this study, no clear rank
order could be found, but they did nd some consistent
dominant or subordinate relationshipswithin that group.
Based on these ndings, they concluded that it now seems
unlikely that interactions between dogs are always, or indeed
ever, driven by the aim to achieve status within a social group.
This conclusion is based on the investigations of a group of
castrated individuals of the same sex, seemingly lacking any
competitive incentives (e.g., females, food or other highly
valued objects). In such a situation, there is not much use to
achieve a higher rank. Furthermore, the concordance between
distributions of behaviors was not investigated before lumping
behaviors into categories. Additionally, formal status signals
like posturing were not included in the analysis as a separate
item. It is a fact that the degree of competition inuences the
performance of conict-related behavior, including utterances
of dominance and submission. So, if competition for resources
is practically absent, and an attenuating inuence on agonistic
behavior might be expected by the neutering, dominance
relationships are expected to be harder to recognize. This also
suggests that generalization from these data is questionable.
The fact that Bradshaw et al. (2009) found some consistent
dominance-related relationships may thus conrm dominance
rather than to deny it.
Being the dominant may in itself become a desired outcome.
This means that conicts about dominance are possible (Bernstein,
1981) because obtaining a higher position may help in securing
resources in the future. Such conicts have been found in other
species also. Andersen et al. (2004) described intense ghting
between piglets that were grouped together and found that the
M.B.H. Schilder et al. / Journal of Veterinary Behavior 9 (2014) 184e191188
intensity was related to the competition level and changes therein
correlated to group size.
7. Learning and the resource holding potential (RHP) model is pro-
posed as alternatives for dominance (Casey, 2008; Bradshaw
et al., 2009). Bradshaw et al. (2009) stated that the develop-
ment of stable relationships can be entirely explained using the
principles of associative learning.Learning, including social
learning, certainly plays an important role in establishing
dominance relationships (Bernstein, 1981; Beacham, 2003).
Bernstein speaks of trained loserswhen individuals after
losing a conict emit submissive behavior toward other indi-
viduals and he denes a dominance relationship as a rela-
tionship in which one can predict that an animal will perform
submissive behavior to a particular animal as a function of a
past history of interactions with that particular animal
(Bernstein, 1981, p. 420).
Spotte (2012, p. 225) also claims that dominance relationships
depend on learning: One animal learns to dominate a familiar
conspecic..However, this nor Bernsteins reasoning does
explain the formal status signaling that instantly may occur
between dogs that meet for the very rst time and also the uni-
directionality in it.
Contrary to the dominance model that predicts relatively stable
relationships to prevent severe future conicts, the RHP model does
not take into account the results of previous interaction. Instead,
ghting ability, motivation, and context play an important role here.
However, the subjective value of a resource may differ between
contestants, and it is this difference that should be a major factor to
predict the outcome of the conict. The asymmetry in ghting
ability is the major aspect of the RHP model (Maynard Smith and
Parker, 1976, p. 159) and in fact includes dominance. So a result
might be that a satiated dominant dog (dominant, because of its
formal signals) may leave food to a subordinate (subordinate,
because of its formal signaling) because its motivation to defend the
asset is low at that specic moment. This example shows that
dominance may function alongside more motivational aspects to
explain an outcome of an interaction, and that both learning and
RHP cannot be seen as alternative explanations for dominance nor
can the incentive value of a resource. Learning should be considered
as a part of the mechanism that enables dominance relationships to
develop, but it leaves the existence of formal dominance/submis-
sion signals unexplained. An individual trait, a tendency, to show
either submissive or dominant behavioral patterns as a coping
strategy cannot be excluded. The RHP model may explain variable
outcomes of conicts (see Barnard and Brown, 1984 for shrews),
whereas formal status signals remain largely independent on the
outcomes of conicts around resources.
Dominance and the humanedog relationship
The 3 recent quantitative studies demonstrate that a limited
number of formal status signals shown by dogs indicate dominance
relationships (Table 1). They conrm the idea of Bauer and Smuts
(2007) that formal dominance in dogs exists. Again other behav-
iors like staring, growling, and showing teeth at the opponent may
primarily signify a tendency to become physically aggressive (a
motivation) rather than having a communicative function con-
cerning status (cf, Fatjo et al., 2007). We can support the notion of
King (2004) and van Kerkhove (2004), that in fact the situation of
dogs in a household may actually more resemble the situation of
wolves in captivity than that of wild wolves or feral dogs. This
would imply that stronger clearer dominance relationships may be
formed or even need to be formed (van Kerkhove, 2004).
Interestingly, extensive literature is available on dominance and
related concepts (status and power) in humans. Social power,
dominance, and submission are considered fundamental di-
mensions of personal relationships (Miller-Day and Jackson, 2012),
pervading every aspect of human social life (Carney et al., 2005).
Much research was done in small groups of unacquainted in-
dividuals, who were gathered to perform some tasks. These persons
were observed and questioned beforehand and/or afterward. It
appeared that a rst assessment of dominance relationships was
made in the rst minute after meeting the other persons in such a
study group, even before any verbal communication had taken place
(Kalma, 1991). Participants of these experiments were able to
indicate who was dominant, but often they were not able to explain
rationally why this was their perception. So, this primary assess-
ment was quick and unconscious. If such a very quick assessment
takes place in our own species where competition and cooperation
are so important, it seems likely that an identical quick assessment
may take place in other social animal species as well. A comparable
quick assessment may occur in dogs and explain the spontaneous
behavioral exchanges between unfamiliar dogs as shortly men-
tioned in the previous section.
McGreevy et al. (2012) give an interesting overview of (dis)
similarities between dogs and people regarding communicative
signals. They mention a number of behaviors (Tables 4, 5, 6: p. 110-
112) like stand over, stare, place paw on to forequarter, averting eye
contact, submissive grin, and submissive posture as signals that
occur in humans and dogs. Apart from the designation submissive
they did not mention that some of these signals could have a
bearing on signaling dominance or submission. Therefore, the
similarities regarding dominance/submission-related communica-
tive signals that we summarize below and in Table 1 are an im-
portant supplementation.
A number of relevant behaviors are reported to be nonverbal
indicators of dominance and submission in humans: sitting in a
straight-up manner (Schwartz et al., 1982) and the chin-high posture
(Zivin, 1977), also called the head tilt or raised head (Mignault and
Chaudhuri, 2003; Carney et al., 2005). On the contrary, if the head
is bowed downward, this turned out to be related to submission
(and some emotions not relevant for this article). A third behavior
mentioned was extended eye contact (Rosa and Masur, 1979; Kalma,
1989), and a fourth behavior was breaking eye contact (Strongman
and Champness, 1968). The latter also may be labeled gaze aver-
sionand is a signal of submissiveness or subordination (Mazur and
Booth, 1998). Gesticulation during speech was mentioned by
Freedman (1972), but it is hard to see how that would be relevant to
dogs. Another nonverbal behavior by dominant people is sitting or
standing on an elevated seat or platform (Eibl-Eibesfeldt, 1997;
Mignault and Chaudhuri, 2003), whereas submissive people make
themselves smaller or take a lower spatial position, for example, by
kneeling, bowing, kissing a hand or shoe, backing down (Morris, 1979;
Mignault and Chaudhuri, 2003).
Reecting on status-related communication signals, it can be
concluded that a number of human characteristics of communica-
tive signals related to dominance and submission resemble distinct
aspects of dogedog communication related to dominance, espe-
cially postural elements. Extended eye contact resembles staring at
an opponent, a signal of an aggressive motivation (Feddersen-
Petersen, 2004) rather than dominance in dogs. The opposite
could be breaking eye contact (e.g., looking away as it is often called
in dog ethology), as a sign of appeasement, one of the means to
avoid (escalation of) a conict (Schenkel, 1967).
These similarities may have facilitated the communication
between both species during domestication. Furthermore, dogs are
likely to interpret human postural information in terms of a domi-
nance/submissive relationship, and they may interpret prolonged
M.B.H. Schilder et al. / Journal of Veterinary Behavior 9 (2014) 184e191 189
eye contact as a threat. Whether humans are able to interpret dog
behavior correctly remains to be seen (see Kujala et al., 2012).
Additionally, it may be expected that dogs may also use a quick
assessment of the dominance position of their human counterpart as
well.Thisinturncouldexplainsomeclinicalobservationsondog
aggression toward humans. It explains why dogs that have an un-
clear rank relationship with their human partner are more likely to
attack when the human partner shows a relatively low posture
(e.g., human lies on the oor, bents down to clean the oor, falls
down, or the dog sits on the humans lap or sits on the couch close to
the human), or when the human is sick or feels weak, or when direct
eye contact is made with a dog (Schilder, 2005). Such kinds of attacks
that may well happen in noncompetitive contexts facilitate the
interpretation that at least some dogs may show a tendency to raise
their status. This is in line with Bernstein (1981) but contrast with the
statement by Bradshaw et al. (2009) that dogs do not tend to do just
that. All these indications suggest that dogs are well able to read at
least an important part of our nonverbal communication relevant to
dominance.
In conclusion, the notion of dominance in dogs in our opinion is
a useful concept. It explains an important aspect of the relationship
between dogs and also between dogs and humans. We want to
emphasize that stability in a relationship does not arise from
exerting dominance (over dogs by humans by using forceful acts)
but by showing formal submissive signals (by the dog toward
humans). Teaching a dog to accept humans as dominants and to
accept related behavioral limits should be different from teaching
appropriate behavior like sitting or lying down on command. The
former depends heavily on learning to accept a submissive status,
which chiey necessitates an adequate socialization of the dog and
a clear and consistent behavior by the owner. The latter involves
reward-based training, which is being facilitated by a clear domi-
nance relationship but not dependent on it. We strongly feel that
this message should be promoted and clearly explained by dog
professionals like instructors, behavior counselors, and veterinar-
ians. Denying the concept of dominance is not helpful to lenient
owners who have a truly dominant dog and are struggling with
control problems. The assessment of the dominance relationship
between dog and owner should be based on observations of pos-
tures and accompanying behaviors shown by the dog. In diagnosing
aggression problems, application of the dominance concept and
observing associated formal status signals in dogs during its in-
teractions in different contexts is very useful, as this gives insight
whether the dogs aggression was fear motivated or whether the
dog was far more condent in its situation and that exerting
dominance in the relationship may play a role. To this purpose, it is
necessary to ask the owners to report formal signals of dominance
or submission of their dog during behavior consultation. Questions
on the postural communication and other (formal) signals the dogs
have shown should always be a part of anamneses. For owners, it is
therefore important to recognize formal signals of submission or
dominance.
Formal signals of active and passive submission are foremost
shown by young dogs toward their owners before puberty. Al-
though testosterone during puberty can boost male dogs to try to
restructure their relationship with their owner, most dogehuman
relationships will overcome this hormone-driven challenge. It is
also very important to realize that aggression is not the equivalent
of dominance; neither is it necessarily an expression of dominant
behavior.
Finally, we want to emphasize that accepting the scientic
concept of dominance should not legitimate the application of forced
dominance signals like alpha rolls in training. They are dangerous in
provoking aggression and will in general not be helpful in estab-
lishing a respectful and harmonious humanedog relationship.
Acknowledgments
The authors are grateful to Frauke Ohl for her helpful comments
on an earlier version to Natalie Cook for correcting the English
language and to 2 referees for useful comments. The idea for the
article was conceived by all 3 authors. The article was mainly
written by the rst author. No specic funding was provided.
Conict of interest
The authors declare no conict of interest.
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... Notwithstanding this claim, the data presented clearly showed that at least some of the dogs formed a linear hierarchy of dominance status. Schilder et al. (2014), commenting on these findings, suggested that a group of human-resourced, sterilized, animals all of the same sex may have had no resources to compete over and thus might not be expected to show much overt social hierarchy. Boitani and Ciucci (1995; see also Van Kerkhove, 2004;Boitani et al., 2007) also suggested that dog groups lack clear hierarchies because they observed multiple breeding individuals -which would not be found in a wolf pack. ...
... I have taken this valuable suggestion further and proposed that dogs' extreme sensitivity to hierarchy in social relationships may be the solution to the apparent paradox of their different behavior toward humans and conspecifics (Wynne, 2021). Several of the formal indicators of dominance and subordinate status in dogs overlap with behaviors used in the same way by humans (Schilder et al., 2014). Thus, van der Borg et al. (2015) identified high posture and muzzle bite as formal dominance indicators in dogs, along with low posture, passing under the head and mouth lick as submission indicators. ...
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Dogs’ remarkable success in living in a human-dominated world rests on a set of adaptations to cohabitation with humans. In this paper, I review the nature of these adaptations. They include changes in reproductive and foraging behavior from their ancestor species, wolves, which can be understood as adaptations to the change from hunting live prey to feeding on human food residues. Dogs also show several changes in social behavior which are more controversial and even somewhat paradoxical. Contrary to theories of canine domestication which view dogs as less aggressive and more cooperative than wolves, several studies show that dogs’ social interactions with conspecifics are more hierarchical and competitive than are wolves’. As scavengers rather than hunters, dogs do not need to cooperate with conspecifics the way that wolves do. But how then can we understand dogs’ willingness to cooperate with humans? I propose an integrated account of dogs’ social behavior that does not assume that dogs need to recognize the species-identity of the individuals with whom they interact. Because of the overlap in formal signals of dominance and submission between dog and human and people’s complete control over the resources dogs need, I propose that people occupy a status of “super-dominance” over dogs. This conception suggests several new lines of research which could shed light on the human-dog relationship to the benefit of both partners.
... Regarding the behavior modification plan (BMP) itself, there is controversy surrounding the appropriateness of the different types of training methods available (simplistically reward versus punishment) (Bradshaw et al., 2009;Schilder et al., 2014;Westgarth, 2016;Todd, 2018). Again, data are lacking in the UK about which training methods are typically employed to resolve unwanted canine behaviors and which category of CP is using them. ...
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Thousands of dogs are relinquished each year in the United Kingdom (UK) owing to behavioral problems. Hence, there is a clear role for behavior modification therapy in the mitigation of this canine welfare risk. Since statutory regulation and a universal register of canine professionals (CPs) does not yet exist in the UK, it remains unclear who is offering such therapy, which behaviors are being treated, what types of approaches various CPs may be taking and the success (or otherwise) CPs are having. This study aimed to provide some insight into these issues from the perspective of pet dog owners (clients). An online survey of 235 participants showed that the primary reason for seeking behavior help from a CP was related to aggression, although often more than 1 issue was reported. Regardless of the behavior problem, no significant differences were found for the type of CP consulted (dog trainer vs. behaviorist). Furthermore, in the client's opinion, there were no significant differences between CP types in their ability to improve their dog's unwanted behavior. Interestingly, behaviorists were significantly more likely than dog trainers to use “reward-based” methods over “balanced training” (balanced being a mix of reward and punishment; x²= 8.226, df = 1, P = 0.004). In conclusion, in the current UK vacuum of statutory regulation, clients are just as likely to employ a trainer as a behaviorist, regardless of their dog's unwanted behavior. However, both CP types were equally able to facilitate behavior improvement, as reported by the client, regardless of the type of training methods (reward-based versus balanced) adopted. This study raises further questions to be explored including the client's opinion of the extent to which the ends (behavior outcomes) justify the means (e.g., punitive training methods which are likely to be aversive for the dog). Also, the general public's understanding of the potential welfare implications of the type of behavior modification plan they may or may not choose to follow.
... "Aggression Misconceptions," a subtheme that was connected to frustration in a lot of participants, covered the perceived tendency of the general public to subscribe to theories that the professional participants disagreed with. Dominance Theory is regularly discussed in dog behavior circles, (Bradshaw, Blackwell, & Casey, 2009;Schilder, Vinke, & Van der Borg, 2014), but was surprisingly sparse in its representation in the professional groups' discussions. This was perhaps because participants had discussed this theory in other fora, leaving little more to be said within the focus groups. ...
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Qualitative methods are increasingly used to investigate the complexities of the dog-human relationship. In order to inform a larger study of human dog interaction, a focus group study was carried out to address the question ‘How is aggressive behavior in dogs perceived and rationalized by people who have experience of dog behavior?’ Six focus groups, including three ‘non-professional’ groups (two groups of dog owners and one group of amateur trainers) and three ‘professional’ groups (a behaviorist group, veterinary group and academic group) were carried out, involving participants who were recruited opportunistically. The focus group transcripts were analyzed using thematic analysis. Findings indicated that participants who do not work with dogs in a professional capacity are largely defensive of dogs when discussing aggressive behavior. However, these participants also discussed factors that make a dog ‘risky’ and how responsible owners manage that risk and the characteristics of ‘dangerous dogs’. For the professional groups, aggressive behavior in dogs presents a barrier to everyday work. They considered working with the owners of dogs showing aggressive behavior and battling anthropogenic stereotypes and misconceptions to be part of the professional challenge. Professionals also contributed views on the nature of ‘dangerous dogs’ and demonstrated awareness of how perceptions could be distorted by the media and propagation of stereotypes. This research highlights the variability of perceptions about canine aggressive behavior. Findings can inform the critical interpretation of quantitative results, and offer a foundation for quantitative study of human directed aggressive behavior in dogs.
... Our research aimed to combine the information received from activity trackers with the subjective assessment and reporting of the owner of the behavior and emotions of the dog. It should be noted that in the dimensional theory of emotions, the dimension of dominance has a separate definition and meaning than in ethology (25). ...
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Leaving a dog home alone is part of everyday life for most dog owners. Previous research shows that dog–owner relationship has multifarious effects on dog behavior. However, little is known about the interplay between dog–owner relationship, physical activity of the dog, and affective experiences at the time of the owner leaving home and reunion when the owner comes home. In this paper, we explored how the general (daily, home alone, and over the 2-week study period) physical activity of the dog, and owner's perceptions of the dog's affective state were correlated at those particular moments. Nineteen volunteer dog owners had their dogs (N = 19) wear two activity trackers (ActiGraph wGT2X-GT and FitBark2) for 2 weeks 24 h/day. Prior to the 2-week continuous physical activity measurement period, the owners filled in questionnaires about the dog–owner relationship and the dog behavior. In daily questionnaires, owners described and assessed their own and their perception of the emotion-related experiences of their dog and behavior of the dog at the moment of separation and reunion. The results indicated that the dog–owner relationship has an interplay with the mean daily and weekly physical activity levels of the dog. An indication of strong emotional dog–owner relationship (especially related to the attentiveness of the dog, continuous companionship, and time spent together when relaxing) correlated positively with the mean daily activity levels of the dog during the first measurement week of the study. Results also suggest that the mean daily and over the 2-week measurement period physical activity of the dog correlated the affective experiences of the dog and owner as reported by the owner when the dog was left home alone. More research is needed to understand the interplay between affect, physical activity of the dog, dog–owner relationship, and the effects of these factors on, and their interplay with, the welfare of dogs.
... Notions of alpha individuals led to dysfunctional and potentially harmful training practices in which humans attempted to dominate domestic dogs. The same notions also led to detrimental labeling of individual dogs as "dominant"-treating dominance like an overarching stable trait-rather than recognizing dominance as a construct specific to particular relationships (Bradshaw et al., 2009(Bradshaw et al., , 2016Mertens, 2004;Schilder et al., 2014;van Kerkhove, 2004). Imposing constructs from human personality models on to other species may lead to similar misattributions and ineffective practices in animal husbandry and welfare. ...
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The Journal of Comparative Psychology has enjoyed a century of publishing some of the best investigations of animal behavior, often with reference to human cognition and behavior. This long history has manifested many paradigm-like shifts. Researchers have fluctuated between treating animals as models of human learning to emphasizing stark differences between animal and human behavior to stressing psychological continuity across species. At this time, there appears little consensus regarding questions of psychological continuity. I argue that this is a futile debate. Rather than focusing on behavior in nonhuman animals that represent potential parallels to human psychology (or behavior), comparative psychologists should focus on questions of development, function, and mechanism of behavior to better understand the behavior of all species in biological context. A focus on understanding underlying mechanisms for behavior rather than settling on behavioral outcomes alone as diagnostic of a species' status on some imaginary scale of progress will help address anthropocentric biases in current approaches. A focus on the "why" and "how" questions espoused by Tinbergen over half a century ago will move the field in better alignment with related fields, such as ethology, and provide greater insights into both animal and human minds. (PsycInfo Database Record (c) 2021 APA, all rights reserved).
Chapter
This chapter explores neurophysiological development and ontogenetic behaviors. The periods of behavioral development and aging are discussed. The ontogeny of attachment, fear reactions, acquisition of self-control, as well as inter- and intraspecific social integration are the subjects of particular attention. The four axes—attachment, fear, self-control, and social integration—constitute the four pillars of development, and, in the course of our work, diseases as well as all behavioral anomalies will be categorized according to these four directions
Chapter
In the current chapter we focus on the social relationships dogs and wolves establish with their pack mates. Dominance and affiliation are relevant features to describe the social relationships of both wolves and dogs. In both species, submissive behaviours and greeting are the best indicators of formal dominance relationships, and in general a linear hierarchy can be detected in most packs. While wolf packs may show a more consistent pattern of clear hierarchical relationships than dogs, subdominant wolves are still more willing to stand their ground to get access to resources, which is accepted by the higher-ranking wolves to some degree. In dogs, this pattern cannot be observed but instead high-ranking animals dominate resources and may escalate into aggression if lower-ranking animals do not keep their distance. In regard to affiliation, measures of proximity and affiliative rates correlate in wolves and dogs, suggesting that both can be used as indexes of bondedness, although, arguably, exchange of affiliative interactions provides a more accurate measure. Behaviours shown in the play contexts in terms of competitive vs. more gentle play reflect the social relationships the animals exhibit outside of play. In accordance, winning positions during dyadic play are exhibited more by the dominant individual in the dyad, and in general there seems little evidence of ‘fair play’ in terms of adherence to the "50:50” rule.
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Research Proposal
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The project is structured as a proposal for the study of relational behavior in dogs (canis familiaris) from a comparative perspective. The work is based on the conceptual framework of Ribes' general psychological theory, in its 2018 update, where relational behavior can be understood as a fundamental characteristic of the so-called comparison contact.
Chapter
This chapter contextualizes the dog-human relationship in the dog's origin as a scavenger on the fringes of human settlements over 15,000 years ago. It then reviews the evidence for unique evolved cognitive structures in dogs that could explain their success in a human-dominated world. Failing to find evidence of unique human-like social-cognitive capacities I then review uncontroversial facts of dogs' basic behavioral biology, including reproductive and foraging behavior and, particularly, affiliative and attachment-related behaviors. This leads to consideration of dogs' social behavior, both conspecific and toward other species, especially humans. I draw attention to a seldom-noted apparent contradiction between dogs' stronger affectional bonds toward humans than toward members of their own species. Dogs' social groups also show steeper social hierarchies accompanied by more behaviors indicating formal dominance than do other canid species including wolves. I resolve this contradiction by proposing that dogs' intense sensitivity to social hierarchy contributes to their willingness to accept human leadership. People commonly control resources that dogs need and also unknowingly express behaviors which dogs perceive as formal signs of dominance. This may be what Darwin was referring to when he endorsed the idea that a dog looks on his master as on a god. Whatever the merits of this idea, if it serves to redirect behavioral research on dogs in human society more toward the social interactions of these species in their diverse forms of symbiosis it will have served a useful function.
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This paper updates and extends Dewsbury's (1982) review of the literature on dominance and reproductive success (RS). The findings from approximately 700 studies are included, over two thirds of which were unavailable to Dewsbury. In order to give a highly condensed and yet meaningful overview, the main findings are represented in four tables, one for male nonprimates, one for female nonprimates, one for male primates, and one for female primates. In the tables for males, findings are analyzed in terms of six different indicators of RS, and in the tables for females, in terms of eight RS indicators.Outside the primate order, evidence largely supported the hypothesis that high-ranking males enjoy greater RS than do subordinate males. For females, studies are more evenly divided between those supporting the hypothesis that high rank and RS are positively correlated and those indicating no significant rank-RS relationship. This may reflect both the lower saliency of hierarchical relationships among females, as well as the lower variability in RS among females, relative to males.Among primates, a complex picture has emerged, especially in the case of males. Much of the complexity appears due to the importance of age and seniority in affecting dominance rank. Also, in some primate species, female preferences for sex partners seem to have little to do with the male's dominance rank, at least at the time mating takes place. Nevertheless, the majority of studies suggest that high- to middle-ranking males have at least a slight lifetime reproductive advantage over the lowest ranking males.
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This dissertation examines dominance, play, affiliation and social preferences in a group of 24 domestic dogs that socialize regularly at a dog daycare facility. In Chapter 1, we analyzed the frequencies and directions of aggression, submission, and dominance displays. The distribution of submission and aggression resulted in a significantly linear hierarchy in the group, and submission was the best indicator of dominance relationships. Age was significantly correlated with dominance rank with older dogs out-ranking younger dogs. Muzzle-licking met most of the criteria for a formal display of submission in dogs, but was displayed in only 18% of relationships. Only 29% of all possible pairs had discernable dominance relationships. In Chapter 2, we examined the relationship between agonism, play and affiliation. Twenty-two percent of all possible pairs had known dominance relationships and also exchanged friendly behaviors (formal relationships), 21% of all pairs exchanged friendly behavior but had no discernable dominance relationship (egalitarian relationships), 50% of all pairs were never observed exchanging friendly or agonistic behaviors (non-interactive relationships), and 7% of pairs had known dominance relationships but did not exchange friendly behavior (agonistic relationships). Friendly behavior was much more frequent than agonistic behavior, and we found a complex association between the two. Egalitarian relationships as well as play and affiliation were more common between males and females than between same-sex pairs. Relationship affinity (an index combining play and affiliation) did not significantly differ in formal versus egalitarian relationships, but the latter were significantly more equitable and playful. In Chapter 3, we investigated patterns of interventions during dyadic play. Interveners directed either (1) a playful behavior at one of the dogs (the target), (2) an affiliative behavior at the target, or rarely, (3) an aggressive behavior at the target. Individual rank did not influence individual interventions rates. Rank relationships between interveners, targets and non-targets did not influence playful interventions. Dogs tended to target higher-ranking dogs during affiliative interventions and target lower-ranking dogs during aggressive interventions, but the latter were too infrequent to apply statistical analyses. Dogs tended to target their preferred partners (“friends”) during play more than support them.
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Wolves are charismatic emblems of wilderness. Dogs, which descended from wolves, are models of urbanity. Do free-ranging dogs revert to pack living or are their societies only reminiscent of a wolfish heritage? Focusing on behavioral ecology, this is the first book to assess societies of both gray wolves and domestic dogs living as urban strays and in the feral state. It provides a comprehensive review of wolf genetics, particularly of New World wolves and their mixture of wolf, coyote and dog genomes. Spotte draws on the latest scientific findings across the specialized fields of genetics, sensory biology, reproductive physiology, space use, foraging ecology and socialization. This interdisciplinary approach provides a solid foundation for a startling and original comparison of the social lives of wolves and free-ranging dogs. Supplementary material, including a full glossary of terms, is available online at www.cambridge.org/9781107015197.
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The evolutionary continuity between humans and other animals suggests that some dimensions of personality may be common across a wide range of species. Unfortunately, there is no unified body of research on animal personality; studies are dispersed across multiple disciplines and diverse journals. To review 19 studies of personality factors in 12 nonhuman species, we used the human Five-Factor Model plus Dominance and Activity as a preliminary framework. Extraversion, Neuroticism, and Agreeableness showed the strongest cross-species generality, followed by Openness; a separate Conscientiousness dimension appeared only in chimpanzees, humans' closest relatives. Cross-species evidence was modest for a separate Dominance dimension but scant for Activity. The comparative approach taken here offers a fresh perspective on human personality and should facilitate hypothesis-driven research on the social and biological bases of personality.