An overview of the dog–human dyad and ethograms within it
ABSTRACT This article reviews the literature on the complex and variable nature of the dog–human dyad and describes the influence of terms such as ''dominance'' on attitudes that humans have toward dogs. It highlights a legacy of tension between ethology and psychology and notes that some practitioners have skills with dogs that elude the best learning theorists. Despite the widespread appeal of being able to communicate with dogs as dogs do with one another, attempting to apply the intraspecific dog etho-gram to human–dog and dog–human interactions may have limited scope. The balance of learning the-ory and ethology on our interactions with dogs is sometimes elusive but should spur the scientific community to examine skills deployed by the most effective humane practitioners. This process will demystify the so-called whispering techniques and permit discourse on the reasons some training and handling techniques are more effective, relevant, and humane than others. This article explores the mismatch between the use of nonverbal communication of 2 species and offers a framework for future studies in this domain. Technologies emerging from equitation science may help to disclose con-fusing interventions through the collar and lead and thus define effective and humane use of negative reinforcement. The case for a validated intraspecific and interspecific canid ethogram is also made.
- SourceAvailable from: Claudia M. Vinke[Show abstract] [Hide abstract]
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, human-dog relationships as well. The concept underlying this model, scientifically defined as an intervening variable reflecting status difference between individuals, is applicable when formal status signals symbolize the long-term relationship between individuals, resulting in a formalized dominance hierarchy. This paper reviews the basics underlying the concept of dominance and reflects on the value and importance of some new quantitative studies on the applicability of the concept of dominance in domestic dogs. The conclusions are, firstly, 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 dog-dog relationships. Secondly, that non-verbal status related communication in humans resembles that in dogs to a considerable degree and hence dogs may be well able to interpret this human status related non-verbal communication from their perspective. Dominance is therefore also likely to play a role in human-dog relationships. Hence, the dominance concept might be useful to explain the development of certain problems in dog-dog and in dog-human relationships. However, enforcing a dominant status by a human may entail considerable risks and should therefore be avoided.Journal of Veterinary Behavior Clinical Applications and Research 07/2014; · 1.65 Impact Factor
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ABSTRACT: Animal training relies heavily on an understanding of species-specific behaviour as it integrates with operant conditioning principles. Following on from recent studies showing that affective states and arousal levels may correlate with behavioural outcomes, we explore the contribution of both affective state and arousal in behavioural responses to operant conditioning. This paper provides a framework for assessing how affective state and arousal may influence the efficacy of operant training methods. It provides a series of three-dimensional conceptual graphs as exemplars to describing putative influences of both affective state and arousal on the likelihood of dogs and horses performing commonly desired behaviours. These graphs are referred to as response landscapes, and they highlight the flexibility available for improving training efficacy and the likely need for different approaches to suit animals in different affective states and at various levels of arousal. Knowledge gaps are discussed and suggestions made for bridging them.Society and Animals 04/2013; 3:300-317. · 0.38 Impact Factor
An overview of the dog–human dyad and ethograms
Paul D. McGreevya, Melissa Starlinga, N. J. Bransonb, Mia L. Cobbc,d, Debbie Calnone
aFaculty of Veterinary Science, University of Sydney, Sydney, NSW, Australia;
bResearch Services Division, Deakin University, VIC, Australia;
cGuide Dogs Victoria, Kew, VIC, Australia;
dAnthrozoology Research Group, Monash University, VIC, Australia; and
eBehaviour Counselling Service, Oakleigh South, VIC, Australia.
Abstract This article reviews the literature on the complex and variable nature of the dog–human dyad
and describes the influence of terms such as ‘‘dominance’’ on attitudes that humans have toward dogs.
It highlights a legacy of tension between ethology and psychology and notes that some practitioners
have skills with dogs that elude the best learning theorists. Despite the widespread appeal of being able
to communicate with dogs as dogs do with one another, attempting to apply the intraspecific dog etho-
gram to human–dog and dog–human interactions may have limited scope. The balance of learning the-
ory and ethology on our interactions with dogs is sometimes elusive but should spur the scientific
community to examine skills deployed by the most effective humane practitioners. This process will
demystify the so-called whispering techniques and permit discourse on the reasons some training
and handling techniques are more effective, relevant, and humane than others. This article explores
the mismatch between the use of nonverbal communication of 2 species and offers a framework for
future studies in this domain. Technologies emerging from equitation science may help to disclose con-
fusing interventions through the collar and lead and thus define effective and humane use of negative
reinforcement. The case for a validated intraspecific and interspecific canid ethogram is also made.
? 2012 Elsevier Inc. All rights reserved.
In the 17th century, English law viewed animals as guilty of
their actions. For example, ‘‘When in 1679 a London
woman swung at Tyburn for bestiality, her canine partner in
crime suffered the same punishment on the same grounds.’’
By the end of the 19th century, the law had changed to view
animals as the property of their human owners, and it
remains this way in many countries today. As science began
to reveal that nature could be subject to human control, the
perception that people were vulnerable to the metaphorical
mystique of animals was rationalized under the banners of
zoology, taxonomy, and veterinary science. The accompa-
nying shift in the nature of the relationship between humans
and animals that remains today is that animals became
objects of human manipulation. For a detailed review of the
social domination of animals in the Victorian era, we refer
the reader to the study by Ritvo (1990).
Since the first half of the 19th century, the sentimental
Address for reprint requests and correspondence: Paul D. McGreevy,
BVSc, MRCVS, PhD, Faculty of Veterinary Science (B19), University of
Sydney, Sydney, NSW 2006, Australia; Tel: 161 2 9351 2810; Fax: 161
2 9351 3957.
1558-7878/$ - see front matter ? 2012 Elsevier Inc. All rights reserved.
Journal of Veterinary Behavior (2012) 7, 103-117
the dog–human relationship is characterized by strong at-
tachment and the optimal well-being of both dogs (Kotrschal
comprehensive review, see Ascione, 2008) and the routine
destruction of unwanted and abandoned dogs (McGreevy
and Bennett, 2010). Canine behavior problems can have an
adverse effect on the well-being of owners and on the wider
community (Voith, 2009), whereas the abandonment or loss
chopathology(Hunt et al., 2008). For these reasons,harmony
between the 2 species is of critical importance.
To better understand canine social behavior and, in turn,
the dog–human dyad, we do well to first study the ways in
which social harmony is created and maintained in stable
groups of dogs. The peacefulness that usually defines such
established dog communities reminds us that there are very
few breaches of social order and that aggression is rare
(Bradshaw et al., 2009). This is underpinned by clear signal-
ing and deference, delivered not demanded (Overall, 1997).
Rough physical contact is far more often part of play than
of violence, and it is foreshadowed by strong signals
This article will review the literature relating to our
current understanding of the complex nature of the dog–
human dyad and examine its characteristics. First, it con-
siders the origins of social-domination belief systems and
social behavior and social learning. Third, it examines
canine–human interactions, examining the extent to which
they reflect or are informed by the canine ethogram and by
learning theory. We summarize by exploring our ability to
apply the canine social ethogram to handling and training
andtoassess the limitations ofthisapproach, andwe suggest
that it is possible to estimate the contribution of ethological
and psychological principles in the manifestations of certain
responses. However, the emphasis here is on estimation, we
are not suggesting that we can quantify the absolute roles of
learned rather than innate responses. Context can determine
whether learning theory will have a greater influence or be
more informative than ethology when training and handling
animals (McGreevy et al., 2009); thus, we avoid using cues
of ethological significance if they run counter to a given
training outcome. For example, because it so reliably trig-
gers playful responses rather than conditioned responses,
play-bowing (an innate canine meta-signal for play) is rarely
used as a visual discriminative stimulus in training. Using
this approach, we offer a framework that helps to describe
this effect in dog–human interactions.
Origins and implications of the social dominance
Social order can be understood as the product of
members of a social group (Petherick, 2010). Social organi-
zation in Canis familiaris can be studied by observing the
way in which dogs gain access to and retain resources
(Drews, 1993). Dominance is characterized as an aspect
of a relationship between 2 or more animals in a social
grouping rather than an attribute or trait of an individual.
The ‘‘dominant’’ animal is considered to have higher status
over another or others in the group (Petherick, 2010). A
dominance relationship involves a simultaneous expression
of both dominance and submission (Schenkel, 1967). A
dominance relationship can be established without any
sign of aggression only indicated by a submissive or
appeasing posture from one of the protagonists. An appeas-
ing attitude from the dog toward humans may therefore
indicate a man–dog dominance relationship. An individual
dog’s motivation to gain access to a particular resource
may be subject to some flux (Bradshaw et al., 2009)
and resource-holding potential may be context-specific
(Shepherd, 2002), but this should not demean its impor-
tance. The ability to learn is similarly context-specific
(McGreevy and Boakes, 2007). It is clear that dogs view
humans differently from the way they view other dogs
(e.g., Rooney et al., 2000). Nevertheless, we would do
well to study the role of intraspecific canine social interac-
tions in dog–human interactions and human–dog interac-
tions, if for no other reason than that humans are
regularly bitten by dogs defending certain resources. Ag-
gression is defined as deliberate threat and/or attack compo-
nents of agonistic behavior with potential to cause injury
associated with conflict and competition (Brain, 2010).
Recent data suggest that higher scores for owner-directed
aggression are associated with male dogs and female
owners (Hsu and Sun, 2010), but this does not necessarily
indicate social dominance as a cause. It is critical that the
role of testosterone in impulsive and reactive aggression
among companion dogs is better understood. And only
when we study how aggression and successful affiliative
activities (such as play) (Horowitz, 2009b) emerge in
dog–dog dyads, will the subtleties of dog body language
and our ability to offend our canine companions, however
inadvertently, become clear.
Social order helps animals within a social group learn
which of them can defend resources and displace another
from them (McGreevy, 2004). This learning is underpinned
by communication and effectively reduces aggression
(McGreevy, 2009). It can exist without the necessity for
individuals to have a sense or concept of their own status.
Humans have gone to great lengths to interpret animal
behavior in terms of social order. Some of the theoretical
constructs that have arisen in the process, such as ‘‘domi-
nance,’’ have acted as obstacles to successfully extending
our understanding of animal behavior (Friedman and
Brinker, 2001). Many authors (e.g., Semyonova, 2003) crit-
icize labels such as ‘‘dominance’’ because they evoke
emotional responses in the observer and can prejudice
interpretations, interfere with verifiability of behavioral
104Journal of Veterinary Behavior, Vol 7, No 2, March/April 2012
responses, and are tightly bound by the human perspective.
It is important to note here that social dominance order (or
orientation) is a term used in psychology to describe a
personality variable that predicts social and political atti-
tudes. It is distasteful to many because it reflects an individ-
ual’s preference for inequality among social groups (Pratto
et al., 1994) and may in some ways relate to right-wing
Buddhism, believe that animals exist in their own right,
rather than as property, as they are represented in Western
law. Thus, it is pertinent to assess the validity of the
dominance construct in terms of Western culture’s sanction
of the control of man over nature (Misra, 1995).
In addition to the effect of cultural attitudes, it is also
worth considering the potential effect of inattentional
blindness on human perception of dog behavior. Inatten-
tional blindness is the notion that ‘‘we rarely see what we
are looking at unless our attention is directed to it’’ (Mack,
2003). This concept highlights the intimate link between
our perception and our attention. Steinker (2007) argued
that by labeling a dog as ‘‘dominant,’’ the humans involved
begin to interpret many behaviors as evidence of ‘‘domi-
nance’’ and ignore any evidence to the contrary. The con-
cept of inattentional blindness may further assist with
understanding why people are very likely to see, remember,
and interpret dog–dog and dog–human interaction in terms
of the dominance framework with which they are familiar.
This concept illustrates the power of our intentions in deter-
mining what we see and what we do not (Mack, 2003).
On a practical level, it is clear that some practitioners
have skills with dogs that elude the best learning theorists;
they may insist that they are using canine ethology and
imposing canine social order to communicate with dogs. As
a result, the role of dog owners and handlers as leaders,
alphas, and trainers is widely debated. A popular view that
has, until recently, prevailed in dog-training circles is that
dog–dog interactions mirror wolf–wolf interactions. An
example is provided by Bauer and Smuts (2007), who
stated that ‘‘as studies of captive wolves found that postural
asymmetries were consistently unidirectional within dyads
and that dominance hierarchies based on these postures
showed a high degree of linearity . such postures were
therefore considered reliable indicators of dominance for
our purposes’’ (in studying companion dogs). These authors
then went on to draw conclusions about play behavior in
companion dogs on the basis of their relative social status,
which were based on wolf data.
Bradshaw et al. (2009) reviewed data available on social
hierarchy in the dog from the perspective of dominance
hierarchy. It seems that, historically, the suggestion that
humans could and should adopt the role of pack leader was
based on 2 implicit assumptions: first, wolves are the ances-
tors of domestic dogs, and second, that a linear hierarchy
the first point is widely accepted, the second has been
frameworks, such as
Semyonova, 2003; van Kerkhove, 2004a; 2004b; Steinker,
cial order as an appropriate model to apply to the social be-
havior of domestic dogs has been widely discussed (van
Kerkhove, 2004a; 2004b; Scott and Fuller, 1965; Serpell,
1995; Bradshaw et al., 2009; Steinker, 2007).
Mech (1999) described 13 years of studying the social
order within wild wolf packs, and the results of this
research contradict most of the widely held beliefs regard-
ing dominance hierarchies in this species that have been
assumed to be applicable to the domestic dog. He found
that family groups, rather than a linear hierarchy, were
observed in wild wolf populations. Based on these results,
van Kerkhove (2004a; 2004b) and Yin (2009) have sug-
gested that ‘‘wolf pack theory’’ does not apply to domestic
dogs and challenged the idea that humans should maintain
social stability in their interactions with dogs by adopting a
‘‘top-dog’’ role. Furthermore, Coppinger (2001) has argued
that dogs are not pack animals. That said, there is evidence
that, where resources are sufficient, large groups of dogs
can occupy a single area with minimal conflict (Bradshaw
et al., 2009) and that dogs can cooperate in tasks
Scott and Fuller (1965) believed that social hierarchies
effectively reduce and minimize overt aggression between
individual members of the pack but noted that a linear hier-
archy does not seem to be a factor for the domestic dog in
maintaining social harmony. They evaluated social order in
their study using a ‘‘dominance test’’ on puppies at 5, 11,
and 15 weeks of age. Two puppies were placed in a pen
with a bone for 10 minutes. Dominance was defined as a
condition in which 1 puppy kept possession of the bone
for at least 8 out of 10 minutes. The dogs were rated as
‘‘dominant,’’ ‘‘incompletely dominant,’’ and ‘‘subordinate,’’
and the effect of dominance on the amount of fighting was
assessed. The observations varied across 3 breeds: Fox
terrier, Basenji, and Shetland sheepdogs. Breed differences
were found in the association between ‘‘dominance’’
(defined as control over a bone) and an effective control
system over fights between individuals. Thus, the notion
that dominance order could control fighting was upheld in
2 of 3 breeds of dogs when observed between 5 and
15 weeks of age. However, if we place these findings in
the context of the difficulty other scientists face when
attempting to develop tests in puppies that predict their
behavior as adults (Wilsson and Sundgren, 1998; Batt
et al., 2009), we can only speculate how any association
between dominance and fighting might vary in dogs of
other ages, let alone in other breeds.
Scott and Fuller (1965) described 9 fundamental behav-
ioral systems for dogs (investigative behavior, epimeletic
behavior, et-epimeletic behavior, allelomimetic behavior,
agonistic behavior, sexual behavior, eliminative behavior,
ingestive behavior, and comfort-seeking behavior [shelter-
seeking]). They concluded that not only were these relevant
(reviewed by [Bradshawet al.,2009;
McGreevy et al Overview of the dog–human dyad 105
to dogs of all breeds, they were also relevant as a frame-
work for comparing human and dog behavior patterns.
While acknowledging that, in many ways, dogs and humans
are different in terms of anatomy, physiology, and behavior,
these authors noted that social behavior patterns are similar
enough, in many contexts, to be mutually recognizable.
A brief consideration of analogues of dog–dog interac-
tions that arise in human–dog dyads suggests that there are
at least some human–dog interactions that align with the
canid intraspecific social ethogram. Analogues, such as
allogrooming, can be useful for humans needing to groom
dogs (McGreevy et al., 2005), whereas others may raise
challenges because there can be tension between ethologi-
cal and psychological constructs in training. For example,
although gaze-averting is a deference display in dogs
(Vas et al., 2005), during some training sessions, it is
critical to keep a dog’s attention focused on the handler.
Furthermore, breed differences in attention to human cues
have been demonstrated (Ga ´csi et al., 2009a), and differ-
ences in aloofness (McGreevy, 2007a) and even retinal
anatomy (McGreevy et al., 2004) may account for the
lack of attention some dogs pay their handlers.
Vas et al. (2005) developed a scale for assessing behav-
ioral responses of dogs to approaches by unfamiliar
humans. These authors note that humans may have selected
dogs based, inter alia, on variation in monitoring of and
response to human cues. Certainly, this could be a first
step in selecting dogs that may excel at protection, guard-
ing, and herding, all of which share related behaviors.
Dogs have been selected for adaptations to human social
life, and these adaptations have led to marked changes in
their communicative, social, cooperative, and attachment
behaviors toward humans. In a review of canine social
cognition, Miklo ´si et al. (2004) state that through a complex
evolutionary process, dogs became adapted for living in
setting now represents a natural ecological niche for this
species. Dogs are extremely flexible in how they process
spatial information and can simultaneously use cues from
different sources and rank the cues based on the complexity
of the environment (Fiset et al., 2000, 2006; Fiset and
LeBlanc, 2007). If humans are part of that environment, we
must consider that dogs trained to work in the presence of
humans who issue discriminative stimuli may disregard
information that they, themselves, collected (Szetei et al.,
2003). Given that companion dogs behaved differently
(attempted a forbidden task) when the owners were in the
room compared with when they were not in the room
(Schwab and Huber, 2006; Horowitz, 2009b), it seems likely
that intimate human–dog relationships (as occur in compan-
ion dogs) may predispose dogs to behave in a socially
dependent manner. All of this suggests that canine scientists
seeking to advance communication in the dog–human dyad
must grant ethologically relevant mechanisms as much
attention as mechanisms that align with learning theory
(McGreevy and Boakes, 2007).
Studying domestic canid social order
Bradshaw et al. (2009) posit that resource holding poten-
tial (Parker, 1974) may be less applicable to dogs than to
other species, but that nevertheless it is useful because it
offers the concept of subjective resource value as a factor
influencing the escalation of conflicts. Bradshaw et al.
(2009) propose that the subjective resource value, in com-
bination with associative learning, explains antagonistic
encounters between dogs more simply than traditional
dominance theory. This is a useful contribution to explain
resource-related aggression but it fails to either acknowl-
edge the possible role of personality dimensions or explain
the mechanisms of dog–dog antagonistic interactions in the
absence of clearly disputed resources.
It could be argued that encounters between 2 members
of a dyad are never resource-neutral. Fighting may be
exhibited in different contexts, including competition over
a resource, and contexts not clearly related to resource
possession (Hahn and Wright, 1998, cited by Wright,
2004). For example, when dogs first meet, one of them is
more the territory holder than the other (even if only by
virtue of being on that site for longer than the protagonist)
and perhaps presence on the territory is a quasi-resource.
But when meeting for the first time, how can dogs be
sure they are valuing or fighting over the same resource?
Before any associative learning about their relationship
can take place, their behavior may represent a manifesta-
tion of positive and negative personality dimensions such
as those described by Sheppard and Mills (2002) or the
5 personality factors described by people interviewed in
the study by Ley et al. (2007).
A given dog’s ability to impose social status at the first
meeting would seem to lay the platform for future interac-
tions over resources. For example, an extroverted dog may
set the stage for subsequent encounters with an introverted
dog over equally valued resources. Some dogs may be
unable to interpret the intentions of unfamiliar dogs when
they approach conspecifics; they may be less concerned
with resources than with the need to obviate a perceived
threat. Clinical experience suggests that some dogs will
behave in an aggressive manner toward any unfamiliar dog,
in any setting, at considerable distances (Debbie Calnon,
personal communication). For these dogs, it is very difficult
to identify the value of a resource because the most likely
primary motivation appears to be fear or anxiety. This
prompts us to ask whether access to self-defense mecha-
nisms (and actions taken to reduce the perceived risk of
harm to self) is the resource these dogs value.
Scientists generally base their measurements of social
hierarchies on who displaces whom from food and, less
often, on who initiates contact with whom. However, it is
important to ask whether the order that predicts displace-
ment and appeasement is sometimes based on relative value
of resources and sometimes on fear. Regardless, the ques-
tion is whether such social order among dogs can include
106Journal of Veterinary Behavior, Vol 7, No 2, March/April 2012
humans and whether perceived breaches of order may
explain how humans occasionally get bitten. It is possible
that inconsistency on the part of humans can create
behavioral conflict in nearby dogs and the resultant frus-
tration can trigger aggression. Dogs that bite humans are
usually, but not always, thought to bite because of fear and
anxiety (McGreevy and Calnon, 2010). Of course, all non-
biting dogs are not necessarily free of fear or anxiety solely
because they are sure of their social status. Nevertheless,
care is warranted in human activities that may amount to
ethologically relevant social threats to dogs.
Conflicts between dogs living in the same household are
most often between members of the same sex, and more
often involve females than males (Sherman et al., 1996;
Wrubel et al., 2011). That said, the triggers for these ag-
gressive encounters are generally reported by dog owners
who are untrained in making behavioral observations.
Thus, it is difficult to identify why these conflicts occur
and why they are more prevalent between members of the
same sex than between members of the opposite sex.
Free-ranging dogs living in groups are reported to show a
linear hierarchy, but although there are differences in fre-
quency of agonistic interactions between males and fe-
males, there is no clear indication whether there is a
separate hierarchy for each sex (Pal et al. 1998). Scott
and Fuller (1965) found that when male and female puppies
contested a resource, males tended to win. They postulated
that this was because males were typically larger. However,
between sexes, size had no effect on the outcome of con-
tests in female–female pairs, and only a weak effect in
male–male pairs. Scott and Fuller (1965) concluded that a
relationship tends to reflect the differential capacities of
the 2 individuals involved. Thus, dogs of different sex
tend to have more defined relationships, with one member
typically playing the dominant role and the other playing
the subordinate, whereas dogs of the same sex tend to
have relationships that are less well-defined, with dominant
and subordinate roles switching readily. The latter relation-
ship is more likely to trigger aggressive behavior generated
The relevance of canine social behavior
Moehlman (1987) reminds us that the regulation of
social structure and behavior in wild canids reflects charac-
teristics of the canid (size, weight, and sex), the group
(group size, territory, and reproductive strategies), and
access to food (temporal and spatial distribution of prey).
Domestication has changed many of these variables beyond
recognition, and perhaps most of all in the supply of
resources, especially food.
Homologous behavioral mechanisms can be identified
between wolves and dogs, but the best model for describing
social relationships among domesticated dogs reared in a
home environment derives from that environment rather
than any wild canid social structure (Wright, 2004). Much
of the work on cognition in dogs has focused on testing pu-
results with captive, usually hand-reared wolves. Wolves in
these populations do not respond to human cues about
location of items in the same way that dogs do, leading to
the conclusion that ‘‘pointing’’ or ‘‘showing’’ are behaviors
developed because of close contact with humans, possibly
because of domestication (Miklosi, 2007).
The ability of dogs to gain access to particular resources,
retain their own resources, and displace other dogs from
resources is a critical element of social order. Dogs are
limited to burying prized objects, but humans can stash
resources in pockets, boxes, and cupboards that only they
can access. Dogs do not feed adult conspecifics nor, for that
matter, do they dictate when they take exercise. A dog’s
behavior can be manipulated by identifying resources
valued by the individual dog. Resources such as food and
exercise can be used by humans to encourage and discour-
age particular canine behaviors, illustrating one distinct
difference between intra- and interspecific social relation-
ships. It follows that resource supply, resource guarding,
and resource-related frustration (on the part of dogs) can
prove problematic in some dog–human dyads.
Most operant conditioning uses learning theory to modify
dog behavior. We control access to the resources and, using
do. Thus, again, we need to be cautious because it may be
access to resources. From ethological descriptors, a social
animal is accorded rank through its ability to gain access to
specific resources. However, we emphasize that it does not
desire to be dominant for the sake of attaining rank per se.
The role of social learning
Social learning occurs when an individual learns by
observation of another individual (Ligout, 2010). In some
species, such as domestic chickens, the social status of
the demonstrator has a strong influence on the perceived
value of the information it imparts (Nicol and Pope,
1999). Social learning is an important aspect of the dog’s
social behavior (Horowitz, 2009a); thus, we should con-
sider how canine ethology informs the way in which we
apply learning theory.
Cognitive tests can include truly novel components that
require learning during the test, including situations in
which dogs learn from watching other dogs successfully
perform and be rewarded for performing a novel task
(Range et al., 2009). It has been shown that the pups of
trained drug-detection bitches learned to pay attention to
target odors from watching their mothers (Slabbert and
Rasa, 1997). In another study, observer dogs were able to
adjust their search behavior for hidden food depending on
McGreevy et alOverview of the dog–human dyad107
the knowledge gained by observing and interacting with a
conspecific (Heberlein and Turner, 2009). It has also been
shown that subordinate dogs, as defined by the balance of
leadership and deferential behavior, learned a detour task
more quickly than dominant dogs if they observed it dem-
onstrated by another dog, but there was no difference if the
demonstrator was human (Pongra ´cz et al., 2008). In a study
of 118 dogs split into 2 groups, the group that watched a
human manipulate a test/treat box was faster in successfully
opening the box and spent more time interacting with it
than the group that did not (Marshall-Pescini et al.,
2008). Thus, we should consider the extent to which dogs
may learn from us with no intended operant conditioning.
The prospect of being able to model behaviors for dogs
to adopt mimicry is beguiling and is strongly hinted at by
studies of model-rival (McKinley and Young, 2003), and
possibly also the effect of pointing and referential gazing
(Elgier et al., 2009; Ga ´csi et al., 2009a,b).
Among the challenges to understanding the way dogs fit
in or not to human households are the variability of dog-
keeping styles (Masters and McGreevy, 2008) and human
personality types (McCrae and John, 1992). Households
can be occupied by single or multiple dogs just as they
can be by single or multiple humans (McGreevy and
In what remains the largest dog behavior research project
of its kind, Scott and Fuller (1965) acknowledged that the
chief relationships they studied were the human males and
females with young dogs. The gender of human participants
we know that dogs’ cortisol concentrations often reduce
with human interaction in general (Hennessy et al., 2001;
Tuber et al., 1996), there is evidence of a more marked
reduction in cortisol responses in shelter dogs when being
stroked by women (Hennessy et al., 1998).
Scott and Fuller (1965) described the complexity of
dog–human interactions and noted that these are variable
and capable of change. They identified 21 possible different
types of social relationships between humans and all the
breeds of dogs. Because several domestic dogs are sexually
neutered, we added the extra categories of neutered male
and neutered female to the original table from Scott and
Fuller (1965). The amended version of this table is repro-
duced later in the text (Table 1) and shows 27 possible
different types of social relationships between dogs and
It seems that, within contemporary households, dogs
may have negligible, frequent, or irregular interactions with
other dogs or people (McGreevy, 2009). Thus, the applica-
bility of canine ethograms may be limited for some dogs
living with humans. Indeed, dogs living in single-dog
households may not even become fluent in their own lan-
guage. Whether this has an effect on their fluency in com-
municating with both conspecifics and heterospecifics is
worth considering. Evidence of the adaptability and flexi-
bility that dogs have when living and communicating
with other species was studied by Feuerstein and Terkel
(2008), who found that a first encounter taking place at
an early age (up to 6 months in cats and up to 1 year in
dogs) enabled most cohabiting dogs and cats to appropri-
ately interpret the particular body language displayed by
the other, even when the signal had an opposite meaning
for both species. Furthermore, they showed that the youn-
ger the animal at first encounter, the better this understand-
ing was, and the more chance there was of establishing an
Social groups in feral dog contexts are arguably subject
to less flux than those in the human–dog domain. Feral
dogs do not meet strangers in their den on a regular basis,
visit parks, or go on holidays. Dogs have not evolved to
know that the new social groups that arise in the company
of strangers, on a visit to a park, or a trip to a holiday
destination, are not going to last for the rest of time.
Fundamental classification of social relationships of dogs and humans
Male (MD) Female (FD)Young (YD)Male (MH) Female (FH) Young (YH)
Male neuter (MN)
Female Neuter (FN)
Adapted from Scott and Fuller, 1965, Genetics and the social behavior of the dog, University of Chicago Press, Chicago, IL. ? 1965 The University of
Chicago. All Rights Reserved.
108 Journal of Veterinary Behavior, Vol 7, No 2, March/April 2012
Making sense of how these novel groupings will affect
access to resources either relies on learning gradually about
each resource and who is allowed it and when, or it
involves some social order, perhaps based entirely on
deference, that removes the need for constant disputes.
There seems to be merit in working out swiftly and
painlessly who must voluntarily defer to whom. Dogs in
the human domain may become highly skilled in respond-
ing appropriately to this cognitive challenge.
Notwithstanding the variability described earlier in the
text, it is possible to map out common dog–human and
human–dog interactions by following the framework for
exploring horse–human interactions offered by McGreevy
et al. (2009). This is discussed later in our concluding
suggestions for further research. However, we need to be
cautious with this approach. Lit et al. (2010) investigated
owner reports of dog–human interactions and showed
how complex owners’ interpretations of such interactions
can be. Elements of the dog–dog social ethogram may look
similar to what humans see when dogs and humans interact
but, so far, we cannot be certain that they look similar
to the dog. We must be aware that dogs may not see or inter-
pret these categories in the same way we do. Nevertheless,
there is merit in offering a framework that has its basis
in dog–dog interactions because it may explain where
errors in human attempts to communicate with dogs most
Using our own observations and pooled experience, and
acknowledging the importance of breed differences in
signaling (Goodwin et al., 1997), we have tabulated the
ways in which dogs interact with each other and with
us, and have contrasted these interactions with some of
the activities we impose on dogs using the following
? Restraint and (giving or receiving) aversive stimuli
? Tactile activities (allogrooming and resting) (Table 3).
? Meeting unfamiliar individuals (Table 4).
? Sharing resources/playing with objects (Table 5).
? Greeting familiar individuals and playing without
objects (Table 6).
? Nontactile interactions (Table 7).
The instances in which dogs do not react to humans as
described in Table 2 are usually because of the relative
difference in height. Common human–dog interactions
in this domain that have no analogue in the canine
ethogram include the use of collars, chains, harnesses,
headcollars, catch poles, lifting dogs off the ground, and
Table 3 shows that dogs will attempt to use almost all
their repertoire of tactile activities with familiar humans.
However, common human–dog interactions that have no
analogue in the canine ethogram include scratching the
chest, putting on collar/harness, combing, brushing, nail-
oral medications, applying topical medications, dressing
wounds, toweling dry, hands-on training, bathing and
cleaning ears, patting as opposed to stroking or scratching,
clipping of coat, kicking, pushing, and smacking.
Table 4 shows that, where our bipedalism does not
interfere, dogs generally attempt to use almost all their
repertoire of greeting activities with familiar humans.
However, our ability to mimic canine responses is severely
limited. Furthermore, common human–dog interactions
that have no analogue in the canine ethogram include hug-
ging familiar group members, patting on head, ignoring,
screaming, and running away.
A consideration of restraint and (giving or receiving) aversive stimuli as they arise in dog–dog, dog–human, and human–dog
Dog–dog behaviorPossible human–dog equivalent (without apparatus) Dog–human analogue of dog–dog behavior
Attempt to escape restraint
Grip with mouth/teeth
Present: hold a body part firmly with hand
Present: lunge forward with hands outstretched
Possible: pin with hands
Possible: pin with hands
Possible: quick lunge with one or both hands
Possible: may do so naturally due to relative height
Pin with chest/body weight
Pin with muzzle
Absentddue to relative difference in height
Absentddue to relative difference in height
Threaten to bite as a form
of body blocking
Absentddue to relative difference in height
aImpeded by human bipedalism.
McGreevy et alOverview of the dog–human dyad109
There are some compelling analogues in Table 5. How-
ever, common human–dog interactions that do not align
with the canine ethogram include throwing objects, not
chasing objects of value (e.g., thrown articles), giving
food (including bones, chews, titbits), giving play objects,
and kicking balls.
Again, Table 6 shows that humans struggle to recipro-
cate in ways that align with the canine ethogram when
playing without objects. In addition, common human–dog
interactions that have no analogue in the canine ethogram
include some of the most popular dog sports: off-lead train-
ing, including agility, canine freestyle, and tricks. It is
worth noting here that, among dogs, it has been found
that more advantaged individuals do not consistently relin-
quish their advantage to facilitate play (Bauer and Smuts,
2007). Role reversals do occur, but certain social conven-
tions influence which behaviors could be used during role
reversals (Bauer and Smuts, 2007).
In Table 7, the imbalance between the 2 species appears
most striking. There are numerous examples of how
humans generally struggle to use elements of the canine
ethogram to communicate with dogs. In addition, there
are common human–dog interactions with no analogue in
the canine ethogram: blowing or whistling in the face, wav-
ing arms around, hand signals, and gesturing (not pointing)
with hands or head.
This examination of intraspecific interactions shows how
commonly dogs seem to deploy elements of their social
ethogram in interactions with humans and where the limits
to these attempts and to reciprocation generally lie. If we
consider contexts in which dogs and humans interact, we
can make predictions about the outcome of the interactions
based on the relative role of learning theory and the canid
intraspecific social ethogram (see Table 8).
Table 8 shows how regularly the most common human
physical interactions (restraint, brushing and combing,
training, feeding, watering, and veterinary care) have no
analogue in the canine social ethogram and how patchy is
the human’s ability to offer plausible analogues of dog–dog
interactions. Taken together, Tables 2-8 show the many
ways in which dogs may attempt to use their social skills
on humans, where their height permits and our bipedalism
does not obstruct such attempts. But they also seem to
demonstrate that, when we consider dog training and
handling, there are limitations to the usefulness of both the
canid ethogram and learning theory. These limitations are
discussed later in the text but, taken together, our shortcom-
ings in the use of the canid ethogram to communicate with
dogs should remind us that successful dog handling is
not successful dog mimicry. However, by the very same
token, we need to acknowledge that these limitations
can help to explain instances of dogs biting humans in
the absence of fear or anxiety. For example, making the
wrong move when a dog has placed its forepaws on
one’s shoulder while staring you in the face can trigger a
bite (McGreevy, 2009).
A consideration of tactile activities (allogrooming and resting) in dog–dog, dog–human, and human–dog interactions
Dog–dog behaviorHuman–dog equivalent Dog–human response?
Resting in physical contact
Present: isolation and restraint of particular body part
Present: massage face/ears with fingers
A consideration of meeting and greeting activities among unfamiliar individuals in dog–dog, dog–human, and human–dog
Dog–dog behavior Human–dog equivalentDog–human response
Increased postural tonus
Lift paw onto forequarter
Sniff under tail
Present: stand tall, tension in body
Present: place hand on shoulder
Present: lunge forward with hands outstretched toward dog
Possible: when lifting a medium-sized dog
Present: not as easily as in dogs.
aImpeded by human bipedalism.
110Journal of Veterinary Behavior, Vol 7, No 2, March/April 2012
Applying the canid ethogram to dog–human
Aspects of our body language and behavior may stim-
ulate resource guarding and different forms of anxiety in
dogs. Human activities that displace a dog may culminate
in a bite if the dog has learned to defend its resources. Dogs
that have been granted free access to some resources as
juveniles learn to defend these and, quite plausibly, others.
Thus, the value of one resource may have implications for
the unexpected defense of others. It is easy to see how this
could lead to unanticipated displays of aggression and to a
dog being labeled dominant.
Our relative height means that dogs automatically look
up to us and that this may lend us an ethological advantage.
It is possible that this makes us ‘‘super-dogs’’: initiators of
expeditions, exercise, grooming, play, and feeding, leaders
who would never be worth questioning, but clearly this is
simplistic. The evidence presented by Bradshaw et al.
(2009) suggests that there is no leader within the dog–dog
domain, so consideration should be given as to how dogs
could transfer such a role to another species. However,
the absence of a clear leader in a study of 19 neutered
male dogs presented by Bradshaw et al. (2009) does not
imply that humans cannot become integrated into social
constructs that have their origins in the social ethology of
Whether dogs have evolved to use their skills for social
order with another species (most notably us) is contentious.
That said, one of the key aspects of evolving with humans
is that dogs have learned to coexist with people, and the
fact that dogs have been so successfully used by people
demonstrates that an effective means of communication
The debate between the relative roles of learning and the
ethogram mirrors the historic tension between ethology and
psychology. On one hand, we have the suggestion that
every response made by a dog (including aggression) has to
be learned and, on the other hand, we have practitioners
proposing that they only have to mimic wolf behavior to
gain compliance and a bond with dogs. The middle ground
suggests that dogs are likely to rely on the social repertoire
they have evolved to use with other dogs unless they have
been socialized with other species.
Communication modalities used in training
In general, we train dogs with negative reinforcement
(NR), positive reinforcement, and punishment. Applying
physical pressure-release (NR) and providing reinforcers
only when a desired behavior is performed (positive
reinforcement) are seldom reported in the canine intra-
specific ethogram, so it seems that we generally train dogs
with modalities that have minimal inherent relevance to
their social learning. This may be a significant failing
on our part.
To be effective and safe in interactions with dogs, our
actions outside operant conditioning rely less on timing and
consistency and more on being able to do the following:
? Interpret dogs’ body language and assess their motiva-
? Mimic appropriate elements of the ethogram (e.g., play-
bowing), notwithstanding their context-specificity;
? Avoid threatening dogs, their young, and their resources;
? Neither inadvertently nor consistently defer and thus
release resources that are valued by the dog;
? Offer appropriate models that may be of use in social
dog–human, and human–dog dyads may share resources
or play with objects
A consideration of the ways in which dog–dog,
Nose objectPresent: push object
A consideration of the ways in which dog–dog, dog–human, and human–dog dyads greet familiar individuals and play
Dog–dog behavior Human–dog equivalentDog–human response
Present: tapping either side of jaw with hands
while dog tries to catch hands
Possible: when lifting a medium-sized dog
Present: reach out with hand
Absent: due to relative
difference in height
Mouth at legs or neck/face
Wrestling while running
aImpeded by human bipedalism.
McGreevy et alOverview of the dog–human dyad111
? Provide useful information (e.g., by pointing at objects
and caches of interest).
These seem to be the very qualities that natural dogfolk
(and possibly many so-called dog whisperers) possess.
Capturing, defining, and measuring these qualities and
training less gifted handlers to reproduce them may be
the enduring legacy of the dog whisperer phenomenon.
Some dogs and some humans seem to have developed
their own fluent means of communication. This involves an
ability to read body language, perhaps underpinned by a
universal interpretation of actions that accompany agonistic
responses in both species: fixed stares, stalking, and high
postural tonus. It may be that dogs learn more quickly when
hand signals are used in training rather than words (Soproni
et al., 2001, 2002). This may support the premise that
handlers who process ideas or communicate pictorially
rather than verbally have made a big leap in their effort
to learn ‘‘dog language,’’ dogs being a visually oriented
(rather than a verbally oriented) species.
in different contexts, but many of the subtleties of canine
ethogram in various common contexts
A conceptual tabulation of the relative roles of learning theory, the value of resources, and the canid intraspecific social
Alignment with the canid
intraspecific social ethogram
Relative role of learning
theory ‘‘How readily can
responses be conditioned?’’
The value of the focal
resource (V), in combination
with associative learning
Human taking resources from dog
Dog taking resources from human
Dog grooming human
Human grooming dog
Hands-off play and training
Hands-on play and training
A consideration of the ways in which dog–dog, dog–human, and human–dog dyads may communicate using nontactile
Dog–dog behavior Human–dog equivalentDog–human response?
Head and neck roll
Relaxed gaze into face
Submissive posture, e.g., inverted
Present: can be imitated with low frequency tone of voice
Present: avert eyes
Present: can be imitated
Possible: can be imitated with higher frequency tone of voice
Possible: can be imitated
Present: can be imitated
Present: can be imitated with high frequency
tones and use of Motherese
Present: when tense or confident
Postural tonus, e.g., U-position, stiff
legs and movement
aImpeded by bipedalism.
112Journal of Veterinary Behavior, Vol 7, No 2, March/April 2012
vocalization remain unknown. Acoustic signals, such as
growls,barks,and whines, havearoleincanine communica-
tion. Growls intended as a warning are of a lower frequency
than growls in play, and are longer, but growls in play have
lower formant frequency dispersions than aggressive
growls (Farago ´ et al., 2010). Formant frequency dispersion
is a measure influenced by the length of the vocal tract;
thus, a low formant frequency dispersion is a reliable
indicator of a larger animal, and a high formant frequency
dispersion indicates the sound is coming from a smaller
animal (Fitch, 1997; Taylor and Reby, 2010). Barks that sig-
nal aggression also have a lower frequency than barks that
signalplay or‘‘happiness,’’andhaveshorter inter-barkinter-
vals (Pongra ´cz et al., 2006). Growls associated with food-
guarding appear to have a strong deterrent effect on other
dogs taking a meaty bone, but warning growls associated
with a threatening stranger are also low in frequency and
have a weaker deterrent effect on dogs taking a meaty bone
(Farago ´ et al., 2010). This would suggest there are elements
of either context or fine acoustic detail that allow dogs
to differentiate between 2 agonistic growls (Farago ´ et al.,
2010). It is possible that dogs interpret low frequency human
vocalizations, including warning tones, as potentially threat-
ening andhighfrequencyhumanvocalizations aspotentially
playful. However, given the likely importance of context in
the interpretation of acoustic signals, it is probable that
the frequency of human vocalizations would be interpreted
considering other cues and the history of the dog with those
and similar human vocalizations. Conversely, it appears
that humans may tend to misinterpret the motivation behind
a canine vocalization as aggressive if it has low frequency
dispersion and thus sounds like a large dog (Taylor et al.,
2010). It is possible there is a perceptual bias in humans to
perceive the growls of large dogs as more aggressive than
growls of small dogs (Taylor et al., 2009).
Humans can alter the types of vocalizations they give
to promote different behavioral responses from dogs.
When humans send a signal consisting of short notes, it
can elicit a reactive response and increase motor activity
levels more than a signal consisting of a longer continuous
note (Yeon, 2007). Dalibard (2009) studied service dogs
through a questionnaire survey and found that voice tone
and clarity affected how well and how quickly dogs
responded to requests.
Coutellier (2006) examined the responses of detection
dogs to the voice commands of their handler or a stranger.
The response of the dogs differed significantly when the
handler and handler’s voice were used, compared with the
stranger and stranger’s voice. There was no difference in
response if only the olfactory cues were changed, indicat-
ing that dogs partially rely on acoustic information to
perform their jobs. Despite the differences in the vocal
range of dogs and humans, there are some similarities
that make some interspecific transfer of information plausi-
ble. For example, it is possible that dogs pick up on the tone
of loudvocalizationsfrom us,allowing them to
discriminate between types of loud vocalizations, such as
those that accompany human anger.
Handlers may also rely on auditory cues from their dogs.
In a study of Hungarian herding dogs (Mudis), listeners,
regardless of their experience with dogs, were able to
categorize bark situations in a way that differed signifi-
cantly from that expected by chance alone (Pongra ´cz et al.,
2005). Associations were strong for particular bark samples
correlated with peak and fundamental frequency and inter-
Relationships between dogs and their handlers
A focus on mechanisms that work in dog training is
useful but may obscure the importance of the relationships
between dogs and their handlers. This possibility has been
studied in military working dog contexts. Lefebvre et al.
(2007) found that the more attention the dog received
from the handler, even if much of this was passive and
simply a result of living and interacting with the handler’s
family, the better its performance and the relationship
between dogs and handlers. Essentially, the more sociable
dogs had better obedience performance than did less socia-
ble dogs. Dogs that spent more time with handlers also
exhibited fewer of the stress-related behaviors that have
been used as indicators of welfare concerns in kenneled
dogs, including pacing, barking, and destruction (Beerda
et al., 1999; Marston et al., 2004).
Horva ´th et al. (2008) studied responses to play sessions
between working dogs and their handlers and concluded
that behaviors associated with control, authority, or aggres-
sion increased cortisol concentrations, whereas play and
affiliative behavior decreased them. There is evidence that
cortisol can act as a hormone-response element and by
doing so can stop the transcription of new proteins neces-
sary to make associative memory, an essential step in
learning (Truss and Beato, 1993). The finding that handler
behavior can be associated with an increase in cortisol con-
centrations has profound implications for working dogs.
Performance of Dutch police dogs trained with and
without shock has been compared (Schilder and van der
Borg, 2004). All dogs in both groups were successful police
dogs, but those in the unshocked group seemed to have a
smoother, more integrated relationship with their handlers,
based on their response to their handlers’ signals.
between obedience and the dog–handler relationship have
been found (Lefebvre et al., 2007). Handlers who interacted
with their dogs in a wider social context (e.g., took their
military working dog home or practiced sports with their
dogs) reported more obedience and fewer bites from
their dogs. The same dogs also showed fewer behaviors
associated with impaired welfare (pacing, barking, destroy-
ing items/materials), indicating that the effects of housing
at a handler’s home and practicing sport were strongly
linked to an enhanced dog–handler relationship.
McGreevy et alOverview of the dog–human dyad 113
Haverbeke et al. (2010) used the Human Familiarization
and Training Program (HFTP) to teach humans to under-
stand normal canine signaling and to reward playful and
appropriate behaviors. When compared with a control
group of dogs not participating in HFTP, the HFTP dogs
carried themselves higher, with less lowering of body pos-
tures associated with fear, showed less yawning, often a
sign of uncertainty, and exhibited fewer aggressive behav-
iors. These authors concluded that because of the changes
in fearfulness, the welfare of HFTP dogs had improved.
The way forward: measuring dog–dog
interactions and human–dog interactions
The social relationship between dogs and humans has
been a topic of great interest in both the popular literature
(e.g., Serpell, 1995; Rogerson, 1988) and, more recently,
scientific literature (e.g., Miklo ´si et al. 2004; Steinker,
2007; Bradshaw et al., 2009). A social relationship may
be defined as regular and predictable behavior occurring
between 2 or more individuals (Scott and Fuller, 1965).
The 2 species interact behaviorally (McConnell, 2002)
and physiologically (Odendaal and Meintjes, 2003), but
empirical studies of these interactions are only just emerg-
ing, despite their profound implications for the success of
individual relationships and the way in which lay people
are educated to manage and train their dogs. Getting
this right can mean the difference between success and
failure in the human–dog interface. Rooney et al. (2000)
showed how intraspecific play styles of individual dogs
were mirrored in those dogs’ play styles with humans.
The complexity of such human–animal interactions is
illustrated by recent studies of the temporal patterning of
human–dog dyadic interactions (Kerepesi et al., 2005). In
a similar vein, Jones and Josephs (2006) reported associa-
tions between salivary hormone concentrations in male
dog handlers and their dogs. Specifically, dogs handled
by men who underwent greater decreases in testosterone
concentration after losing a dog agility competition showed
greater increases in cortisol concentrations than in winning
teams (Jones and Josephs, 2006).
Although defining social relationships as regular and
predictable behavior occurring between 2 or more individ-
uals, Scott and Fuller (1965) also noted that they are neither
invariable nor incapable of change. Although we agree with
ship (changing with factors such as ontogeny, context, etc.),
we propose that quantifying certain features of the
dog–human interaction may provide further insights that
increase our understanding of why some dog–human rela-
tionships are successful and others are not. Management
factors have a significant impact on the success rate of dogs
learning particular tasks (Batt et al., 2010) and in facilitating
to train a dog to walk next to a handler may provide insights
dog. In this context, ‘‘communicate effectively’’ is meant to
refer to the handler’s ability to train the dog to do the desired
task. For example, a dog can be trained to maintain a consis-
tent position relative to the handler when walking on the
lead by strategically releasing the lead pressure when the
dogisinthecorrectposition.Itmay also reflect subtle exam-
ples of best practice in the way handler’s posture changes
during heelwork (McGreevy, 2009).
Thus, here we see some exciting parallels between the
science of dog training and equitation science––the science
of horse riding and training (McGreevy, 2007b). Capturing,
defining, and measuring the qualities of techniques used by
the very best dog handlers are the essence of this approach.
It seems that, despite the importance of on- and off-lead
heelwork in the eyes of seasoned trainers, many owners use
the lead simply to restrain the dog and many dogs use the
lead to control their owners. That said, we should not ignore
this critical interface. Even outside formal training, the aver-
age pet dog spends at least some time on the lead. Such
periods are likely to be of enormous relevance to the dog be-
the endpoint (such as arrival at an exercise venue) is often
highly reinforcing. In equitation science, rein tension meters
are an established means of measuring the mechanisms
horse riders use to train horses with NR (i.e., the horse is
rewarded by the rider immediately releasing the tension in
the rein when a horse performs the desired behavior)
(McGreevy and McLean, 2007). Using the same approach,
alead tension meter could be used torecord thegeneral level
of contact, the intensity and frequency of all corrections a
handler applies through the lead, and the dog’s latency to re-
spond to them. Such data would capture the effectiveness
to which the dog has become habituated to aversive stimuli
from the collar.
Considering principles beyond NR, equitation science
shows how operant conditioning of pressure cues can lay
the foundation for the emergence of classical conditioned
cues such as those from the rider’s seat or, most relevant
here, the handler’s posture. It explains how even without
leading a horse or applying physical pressure, hands-off
round pen techniques are effective in training horses. As
such, it identifies some fascinating commonalities with the
use of body language by dog trainers. The play-bow and the
body block (McConnell, 2002) are examples of postural
techniques that seasoned dogfolk use, often without being
aware of what they are doing. We can use kinematic anal-
ysis systems that are emerging from equitation science to
study these techniques.
Equitation science has identified the need for a working
horse ethogram (Heleski et al., 2009). By the same token,
canine scientists need a validated canid intraspecific social
ethogram that exhaustively lists the types of behavior per-
formed by domestic dogs in a social context (Plowman,
114Journal of Veterinary Behavior, Vol 7, No 2, March/April 2012
2010). It is accepted that many research groups have devel-
oped their own ethograms but recent meetings of canine
scientists (such as the first and second Canine Science
Forum in 2008 and 2010, respectively) serve to emphasize
the need for a canine ethogram that is validated to with-
stand use by multiple observers and different laboratories
around the world. Lack of standardization will continue
to suppress the development of clear communication among
groups and obstruct attempts at meta-analysis (Tomkins
et al., 2010). A validated ethogram will pave the way for
the development of a related and cross-referenced dog–
human ethogram. This tool is of critical importance because
with it canine scientists can advise veterinarians, handlers,
trainers, and owners on the relative importance of subtle
behavioral mechanisms operating at the dog–human inter-
face. Only then will we truly decipher the role, if any, of
perceived social status in our dogs’ responses.
It is critical that we study the way in which harmony is
achieved among dogs and the extent to which communica-
tion between dogs and humans can align with or breach the
framework offered by the dog–dog social ethogram. As-
pects of our body language and behavior may stimulate
resource-guarding and forms of anxiety in dogs. Displacing
dogs that have learned to defend their resources may
culminate in a bite. It is critical that we fully explore
possible mechanisms beyond a learning-theory framework
that may explain dog–human aggression and that the
emerging science of dog training continues to embrace
principles derived from both psychology and ethology.
The authors thank Dr Wendy Birkhead for enlightening
discussions on the problems faced by anthropologists when
grappling with the concept of ‘‘family.’’ Two anonymous
reviewers provided excellent feedback on earlier drafts of
Ascione, F., 2008. The international handbook of animal abuse and
cruelty: theory, research and application. Purdue University Press,
West Lafayette, IN.
Brain, P.F., 2010. Aggression. In: Mills, D.S., Marchant-Forde, J.N.,
McGreevy, P.D., Morton, D.B., Nicol, C.J., Phillips, C.J., Sandoe, P.,
Swaisgood, R.R. (Eds.), Encyclopedia of Applied Animal Behaviour
and Welfare. CABI Publishing, Wallingford, Oxon, pp. 8-10.
McCabe, H.M., 2003. Benefits of interacting with companion animals.
Am. Behav. Sci. 47, 94-99.
Batt, L., Batt, M., Baguley, J., McGreevy, P.D., 2009. The value of puppy
raisers’ assessments of potential guide dogs’ behavioral tendencies
and ability to graduate. Anthrozo ¨os 22, 71-76.
J.W., Karpf,A.S., Suthers-
Batt, L., Batt, M., Baguley, J., McGreevy, P.D., 2010. Relationships
between puppy management practices and reported measures of suc-
cess in guide dog training. J. Vet. Behav.: Clin. Appl. Res. 5, 240-246.
Bauer, E.B., Smuts, B.B., 2007. Cooperation and competition during
dyadic play in domestic dogs, Canis familiaris. Anim. Behav. 73,
Beerda, B., Schilder, M.B., Bernadina, W., van Hooff, J.A., de Vries, H.W.,
Mol, J.A., 1999. Chronic stress in dogs subjected to social and spatial
restriction: part II. Hormonal and immunological responses. Physiol.
Behav. 66, 243-254.
Bradshaw, J.W., Blackwell, E.J., Casey, R.A., 2009. Dominance in domes-
tic dogsduseful construct or bad habit? J. Vet. Behav.: Clin. Appl.
Res. 4, 135-144.
Coppinger, R., 2001. Dogs: A Startling New Understanding of Canine Or-
igin, Behavior and Evolution. Scribner, New York, NY.
Coutellier, L., 2006. Are dogs able to recognize their handler’s voice? A
preliminary study. Anthrozo ¨os 19, 278-284.
Dalibard, G.H., 2009. Parameters influencing service dogs’ quality of
response to commands: retrospective study of 71 dogs. J. Vet. Behav.:
Clin. Appl. Res. 4, 19-24.
Drews, C., 1993. The concept and definition of dominance in animal
behaviour. Behaviour 125, 283-313.
Elgier, A.M., Jakovcevic, A., Barrera, G., Mustaca, A.E., Bentosela, M.,
2009. Communication between domestic dogs (Canis familiaris) and
humans: dogs are good learners. Behav. Processes 81, 402-408.
Farago ´, T., Pongra ´cz, P., Range, F., Vira ´nyi, Z., Miklo ´si, A´., 2010. ‘The
bone is mine’: affective and referential aspects of dog growls.
Anim. Behav. 79, 917-925.
Feuerstein, N., Terkel, J., 2008. Interrelationships of dogs (Canis
familiaris) and cats (Felis catus L.) living under the same roof.
Appl. Anim. Behav. Sci. 113, 150-165.
Fiset, S., Gagnon, S., Beaulieu, C., 2000. Spatial encoding of hidden
objects in dogs (Canis familiaris). J. Comp. Psychol. 114, 315-324.
Fiset, S., Landry, F., Ouellette, M., 2006. Egocentric search for disappear-
ing objects in domestic dogs: evidence for a geometric hypothesis of
direction. Anim. Cogn. 9, 1-12.
Fiset, S., LeBlanc, V., 2007. Invisible displacement understanding in
domestic dogs (Canis familiaris): the role of visual cues in search
behavior. Anim. Cogn. 10, 211-224.
Fitch, W.T., 1997. Vocal tract length and formant frequency dispersion
correlate with body size in rhesus macaques. J. Acoust. Soc. Am.
Friedman, S.G., Brinker, B., 2001. The struggle for dominance: fact or
fiction? A bird’s eye view. Original Flying Machine 17-20.
Ga ´csi, M., McGreevy, P., Kara, E., Miklo ´si, A´, 2009a. Effects of selection
for cooperation and attention in dogs. Behav. Brain Funct. 5, 31.
Ga ´csi, M., Gyoo ¨ri, B., Vira ´nyi, Z., Kubinyi, E., Range, F., Bele ´nyi, B.,
Miklo ´si, A´, 2009b. Explaining dog wolf differences in utilizing human
pointing gestures: selection for synergistic shifts in the development of
some social skills. PLoS One 4, e6584.
Goodwin, D., Bradshaw, J.W., Wickens, S.M., 1997. Paedomorphosis
affects agonistic visual signals of domestic dog. Anim. Behav. 53,
Hahn, M.E., Wright, J.C., 1998. The influence of genes on social behavior
of dogs. In: Grandin, T. (Ed.), Genetics and the Behavior of Domestic
Animals. Academic, New York, NY, pp. 299-318.
Haverbeke, A., Rzepa, C., Depiereux, E., Deroo, J., Giffroy, J.M.,
Diederich, C., 2010. Assessing efficiency of a Human Familiarisation
and Training Programme on fearfulness and aggressiveness of military
dogs. Appl. Anim. Behav. Sci. 123, 143-149.
Heberlein, M., Turner, D.C., 2009. Dogs, Canis familiaris, find hidden
food by observing and interacting with a conspecific. Anim. Behav.
Heleski, C.R., McGreevy, P.D., Kaiser, L.J., Lavagnino, M., Tans, E.,
Bello, N., Clayton, H.M., 2009. Effects on behaviour and rein tension
on horses ridden with/without martingales and rein inserts. Vet. J. 181,
McGreevy et alOverview of the dog–human dyad115
Hennessy, M.B., Voith, V.L., Mazzei, S.J., Buttram, J., Miller, D.D.,
shelter, and an exploration of the ability of these measures to predict
problem behavior after adoption. Appl. Anim. Behav. Sci. 73, 217-233.
Hennessy, M.B., Williams, M.T., Miller, D.D., Douglas, C.W., Voith, V.L.,
1998. Influence of male and female petters on plasma cortisol and be-
haviour: can human interaction reduce the stress of dogs in a public
animal shelter? Appl. Anim. Behav. Sci. 61, 63-77.
Horowitz, A.C., 2009a. Attention to attention in domestic dog (Canis
familiaris) dyadic play. Anim. Cogn. 12, 107-118.
Horowitz, A., 2009b. Disambiguating the "guilty look": Salient prompts to
a familiar dog behaviour. Behav. Processes 81, 447-452.
Horva ´th, Z., Do ´ka, A., Miklo ´si, A´, 2008. Affiliative and disciplinary
behavior of human handlers during play with their dog affects cortisol
concentrations in opposite directions. Horm. Behav. 54, 107-114.
Hsu, Y., Sun, L., 2010. Factors associated with aggressive responses in pet
dogs. Appl. Anim. Behav. Sci. 123, 108-123.
Hunt, M., Al-Awadi, H., Johnson, M., 2008. Psychological sequelae of pet
loss following Hurricane Katrina. Anthrozo ¨os 21, 109-121.
Jones, A.C., Josephs, R.A., 2006. Interspecies hormonal interactions
between man and the domestic dog (Canis familiaris). Horm. Behav.
Kerepesi, A., Jonsson, G.K., Miklo ´si, A´, Topa ´l, J., Csa ´nyi, V.,
Magnusson, M.S., 2005. Detection of temporal patterns in dog–human
interaction. Behav. Processes 70, 69-79.
Kotrschal, K., Scho ¨berl, I., Bauer, B., Thibeaut, A., Wedl, M., 2009.
Dyadic relationships and operational performance of male and female
owners and their male dogs. Behav. Processes 81, 383-391.
Lefebvre, D., Diederich, C., Delcourt, M., Giffroy, J.M., 2007. The quality
of the relation between handler and military dogs influences efficiency
and welfare of dogs. Appl. Anim. Behav. Sci. 104, 49-60.
Ley, J., Bennett, P.C., Coleman, G.J., 2007. Personality dimensions that
emerge in companion canines. Appl. Anim. Behav. Sci. 110, 305-317.
Ligout, S., 2010. Social learning. In: Mills, D.S., Marchant-Forde, J.N.,
McGreevy, P.D., Morton, D.B., Nicol, C.J., Phillips, C.J., Sandoe, P.,
Swaisgood, R.R. (Eds.), Encyclopedia of Applied Animal Behaviour
and Welfare. CABI Publishing, Wallingford, Oxon, p. 563.
Lit, L., Schweitzer, J.B., Oberbauer, A.M., 2010. Characterization of
human–dog social interaction using owner report. Behav. Processes
Mack, A., 2003. Inattentional blindness: looking without seeing. Curr. Dir.
Psychol. Sci. 12, 180-184.
Marshall-Pescini, S., Valsecchi, P., Petak, I., Accorsi, P., Previde, E., 2008.
Does training make you smarter? The effects of training on dogs’
performance (Canis familiaris) in a problem-solving task. Behav.
Processes 78, 449-454.
Marston, L.C., Bennett, P.C., Coleman, G.J., 2004. What happens to
shelter dogs? An analysis of data for 1 year from three Australian
shelters. J. Appl. Anim. Welf. Sci. 7, 27-47.
Masters, A., McGreevy, P., 2008. Dog keeping practices as reported by
McConnell, P.B., 2002. The Other End of the Leash: Why We Do What
We Do Around Dogs. Bantam, Milsons Point, NSW.
McCrae, R.R., John, O.P., 1992. An introduction to the five-factor model
and its applications. J. Pers. 60, 175-215.
McGreevy, P.D., 2004. Equine Behavior–A Guide for Veterinarians and
Equine Scientists. W.B. Saunders, London.
McGreevy, P.D., 2007a. Breeding for quality of life. Anim. Welf. 16,
McGreevy, P.D., 2007b. The advent of equitation science. Vet. J. 174,
McGreevy, P.D., 2009. A modern dog’s life. UNSW Press, Randwick, NSW.
McGreevy, P.D., Bennett, P.B., 2010. Challenges and paradoxes in the
companion animal niche. Darwinian selection, selective breeding
and the welfare of animals. Anim. Welf. 19, 11-16.
McGreevy, P.D., Boakes, R.A., 2007. Carrots and Sticks: The Principles of
Animal Training. Cambrdige University Press, Cambridge, UK.
McGreevy, P.D., Calnon, D., 2010. Canine aggression in perspective. Vet.
J. 186, 1-2.
McGreevy, P.D., Grassi, T.D., Harman, A.M., 2004. A strong correlation
exists between the distribution of retinal ganglion cells and nose
length in the dog. Brain Behav. Evol. 63, 13-22.
McGreevy, P.D., Masters, A.M., 2008. Risk factors for separation related
distress and feed-related aggression in dogs: additional findings
from a survey of Australian dog owners. Appl. Anim. Behav. Sci.
McGreevy, P.D., McLean, A.N., 2007. The roles of learning theory and
ethology in equitation. J. Vet. Behav.: Clin. Appl. Res. 2, 108-118.
McGreevy, P., Oddie, C., Burton, F., McLean, A., 2009. The horse-human
dyad: can we align horse training and handling activities with the
equid social ethogram? Vet. J. 181, 12-18.
McGreevy, P.D., Righetti, J., Thomson, P.C., 2005. The reinforcing value
of physical contact and the effect on canine heart rate of grooming
in different anatomical areas. Anthrozo ¨os 18, 236-244.
McKinley, S., Young, R.J., 2003. The efficacy of the model-rival method
when compared with operant conditioning for training domestic
dogs to perform a retrieval-selection task. Appl. Anim. Behav. Sci.
Mech, L.D., 1999. Alpha status, dominance, and division of labor in wolf
packs. Canad. J. Zool. 77, 1196-1203.
Miklo ´si, A´, 2007. Big thoughts in small brains? Dogs as a model for
understanding human social cognition. Cogn. Neurosci. Neuropsy-
chol. 18, 467-471.
Miklo ´si, A´, Topa ´l, J., Csa ´nyi, V., 2004. Comparative social cognition: what
can dogs teach us? Anim. Behav. 67, 995-1004.
Misra, R.P., 1995. Environmental Ethics: A Dialogue of Cultures. Concept
Publishing Company, New Delhi, India, p. 103.
Moehlman, P.D., 1987. Social organization in jackals. Am. Sci. 75,
Nicol, C., Pope, S., 1999. The effects of demonstrator social status and
prior foraging success on social learning in laying hens. Anim. Behav.
Odendaal, J.S., Meintjes, R.A., 2003. Neurophysiological correlates of
affiliative behaviour between humans and dogs. Vet. J. 165, 296-301.
Overall, K.L., 1997. Clinical Behavioral Medicine for Small Animals,
Mosby, St. Louis, MO, pp. 11, 28.
Pal, S., Ghosh, B., Roy, S., 1998. Agonistic behaviour of free-ranging dogs
(Canis familiaris) in relation to season, sex and age. Appl. Anim.
Behav. Sci. 59, 331-348.
Parker, G.A., 1974. Assessment strategy and the evolution of animal con-
flicts. J. Theor. Biol. 47, 223-243.
Petherick, J.C., 2010. Dominance. In: Mills, D.S., Marchant-Forde, J.N.,
McGreevy, P.D., Morton, D.B., Nicol, C.J., Phillips, C.J., Sandoe, P.,
Swaisgood, R.R. (Eds.), Encyclopedia of Applied Animal Behaviour
and Welfare. CABI Publishing, Wallingford, Oxon, pp. 185-186.
Plowman, A., 2010. Ethogram. In: Mills, D.S., Marchant-Forde, J.N.,
Sandoe, P., Swaisgood, R.R. (Eds.), Encyclopedia of Applied Animal
Behaviour and Welfare. CABI Publishing, Wallingford, Oxon,
Pongra ´cz, P., Molna ´r, C., Miklo ´si, A´, Csa ´nyi, V., 2005. Human listeners are
able to classify dog (Canis familiaris) barks recorded in different
situations. J. Comp. Psychol. 119, 136-144.
Pongra ´cz, P., Molna ´r, C., Miklo ´si, A´, 2006. Acoustic parameters of dog
barks carry emotional information for humans. Appl. Anim. Behav.
Sci. 100, 228-240.
Pongra ´cz, P., Vida, V., Ba ´nhegyi, P., Miklo ´si, A´, 2008. How does domi-
nance rank status affect individual and social learning performance
in the dog (Canis familiaris)? Anim. Cogn. 11, 75-82.
Pratto, F., Sidanius, J., Stallworth, L.M., Malle, B.F., 1994. Social domi-
nance orientation: a personality variable predicting social and political
attitudes. J. Pers. Soc. Psychol. 67, 741-763.
Range, F., Heucke, S.L., Gruber, C., Konz, A., Huber, L., Vira ´nyi, Z.,
2009. The effect of ostensive cues on dogs’ performance in a
116Journal of Veterinary Behavior, Vol 7, No 2, March/April 2012
manipulative social learning task. Appl. Anim. Behav. Sci. 120,
Ritvo, H., 1990. The Animal Estate. Penguin Books, New York, NY.
Rogerson, J., 1988. Your Dog: Its Development, Behaviour and Training.
Popular Dogs Publishing, London.
Rooney, N.J., Bradshaw, J.W., Robinson, I.H., 2000. A comparison of
dog–dog and dog–human play behaviour. Appl. Anim. Behav. Sci.
Schenkel, R., 1967. Submission; its features and function in the wolf and
dog. Am. Zool. 7, 319-329.
Schilder, M.B., van der Borg, J.A., 2004. Training dogs with help of the
shock collar: short and long-term behavioural effects. Appl. Anim.
Behav. Sci. 85, 319-334.
Schwab, C., Huber, L., 2006. Obey or not obey? Dogs (Canis familiaris)
behave differently in response to attentional states of their owners.
J. Comp. Psychol. 120, 169-175.
Scott, J., Fuller, J., 1965. Genetics and the Social Behavior of the Dog.
University of Chicago Press, Chicago, IL.
Semyonova, A., 2003. The social organization of the domestic dog; a
longitudinal study of domestic canine behavior and the ontogeny of
domestic canine social systems. The Carriage House Foundation, The
Hague, Netherland. Version 2006. Available at: www.nonlineardogs.
com. Accessed October 15, 2010.
Serpell, J., 1995. The Domestic Dog: I Evolution, Behaviour, and Interac-
tions With People. Cambridge University Press, Cambridge, UK,
New York, NY.
Shepherd, K., 2002. Development of behaviour, social behaviour and com-
munication in dogs. In: Horwitz, D.F., Mills, D.S., Heath, S. (Eds.),
BSAVA Manual of Canine and Feline Behavioural Medicine. BSAVA,
Quedgeley, UK, pp. 8-20.
Sheppard, G., Mills, D.S., 2002. The development of a psychometric scale
for the evaluation of the emotional predispositions of pet dogs. Int.
J. Comp. Psychol. 15, 201-222.
Sherman, C., Reisner, I., Taliaferro, L., Houpt, K., 1996. Characteristics,
treatment, and outcome of 99 cases of aggression between dogs.
Appl. Anim. Behav. Sci. 47, 91-108.
Slabbert, J.M., Rasa, O.A., 1997. Observational learning of an acquired
maternal behaviour pattern by working dog pups: an alternative train-
ing method? Appl. Anim. Behav. Sci. 53, 309-316.
Soproni, K., Miklo ´si, A´, Topa ´l, J., Csa ´nyi, V., 2001. Comprehension of
human communicative signs in pet dogs (Canis familiaris). J. Comp.
Psychol. 115, 122-126.
Soproni, K., Miklo ´si, A´, Topa ´l, J., Csa ´nyi, V., 2002. Dogs’ (Canis
familiaris) responsiveness to human pointing gestures. J. Comp.
Psychol. 116, 27-34.
Steinker, A., 2007. Terminology Think Tank: Social dominance theory as
it relates to dogs. J. Vet. Behav.: Clin. Appl. Res. 2, 137-140.
Szetei, V., Miklo ´si, A´, Topa ´l, J., Csa ´nyi, V., 2003. When dogs seem to lose
their nose: an investigation on the use of visual and olfactory cues in
communicative context between dog and owner. Appl. Anim. Behav.
Sci. 83, 141-152.
Taylor, A.M., Reby, D., McComb, K., 2009. Context-related variation in
the vocal growling behaviour of domestic dogs, Canis familiaris.
Ethology 115, 905-915.
Taylor, A., Reby, D., McComb, K., 2010. Why do large dogs sound more
aggressive to human listeners: acoustic bases of motivational misattri-
butions. Ethology 116, 1155-1162.
Taylor, A.M., Reby, D., 2010. The contribution of source-filter theory to
mammal vocal communication research. J. Zool. 280, 221-236.
Tomkins, L.M., McGreevy, P.D., Branson, N.J., 2010. Lack of standardiza-
tion in reporting motor laterality in the dog. J. Vet. Behav.: Clin. Appl.
Res. 5, 235-239.
Truss, M., Beato, M., 1993. Steroid hormone receptors: interaction with
deoxyribonucleic acid and transcription factors. Endocr. Rev. 14,
Tuber, D.S., Hennessy, M.B., Sanders, S., Miller, J.A., 1996. Behavioral
and glucocorticoid responses of adult domestic dogs (Canis familiaris)
van Kerkhove, W., 2004a. A fresh look at the wolf-pack theory of
companion-animal dog social behavior. J. Appl. Anim. Welf. Sci. 7,
van Kerkhove, W., 2004b. Response to comments. J. Appl. Anim. Welf.
Sci. 7, 299-300.
Vas, J., Topa ´l, J., Ga ´csi, M., Miklo ´si, A´, Csa ´nyi, V., 2005. A friend or an
enemy? Dogs’ reaction to an unfamiliar person showing behavioural
cues of threat and friendliness at different times. Appl. Anim. Behav.
Sci. 94, 99-115.
Voith, V.L., 2009. The impact of companion animal problems on society
and the role of veterinarians. Vet. Clin. North Am. Small Anim. Pract.
Wilsson, E., Sundgren, P.E., 1998. Behaviour test for eight-week-old
puppies-heritabilities of tested behaviour traits and its correspondence
to later behaviour. Appl. Anim. Behav. Sci. 58, 151-162.
Wright, J.C., 2004. Comment on van Kerkhove’s "Wolf-Pack Theory".
J. Appl. Anim. Welf. Sci. 7, 295-298.
Wrubel, K.M., Moon-Fanelli, A., Maranda, L., Dodman, N., 2011. Inter-
dog household aggression: 38 cases (2006–2007). J. Am. Vet. Med.
Assoc. 238, 731-740.
Yeon, S.C., 2007. The vocal communication of canines. J. Vet. Behav.:
Clin. Appl. Res. 2, 141-144.
Yin, S., 2002. A new perspective on barking in dogs (Canis familiaris).
J. Comp. Psychol. 116, 189-193.
Yin, S., 2009. Low Stress Handling, Restraint and Behaviour Modification
of Dogs and Cats. Cattle Dog Publishing, Davis, CA.
McGreevy et alOverview of the dog–human dyad117