Trainability and boldness traits differ between dog breed clusters based on conventional breed categories and genetic relatedness

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DOI: 10.1016/j.applanim.2011.03.006
Cite this publication
Modern dog breeding has given rise to more than 400 breeds differing both in morphology and behaviour. Traditionally, kennel clubs have utilized an artificial category system based on the morphological similarity and historical function of each dog breed. Behavioural comparisons at the breed-group level produced ambiguous results as to whether the historical function still has an influence on the breed-typical behaviour. Recent genetic studies have uncovered genetic relatedness between dog breeds, which can be independent from their historical function and may offer an alternative explanation of behavioural differences among breeds. This exploratory study aimed to investigate the behaviour profiles of 98 breeds, and the behavioural differences among conventional breed groups based on historical utility and among genetic breed clusters. Owners of 5733 dogs (98 breeds) filled out an online questionnaire in German. Breed trait scores on trainability, boldness, calmness and dog sociability were calculated by averaging the scores of all individuals of the breed. Breeds were ranked on the four traits and a cluster analysis was performed to explore behavioural similarity between breeds.
Applied Animal Behaviour Science 132 (2011) 61–70
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Applied Animal Behaviour Science
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Trainability and boldness traits differ between dog breed clusters
based on conventional breed categories and genetic relatedness
Borbála Turcsán, Enik ˝
o Kubinyi, Ádám Miklósi
Department of Ethology, Eötvös University, H-1117, Pázmány P. s. 1/c, Budapest, Hungary
article info
Article history:
Accepted 2 March 2011
Available online 31 March 2011
Breed characteristics
Typical behaviour
Genetic relatedness
Modern dog breeding has given rise to more than 400 breeds differing both in morphol-
ogy and behaviour. Traditionally, kennel clubs have utilized an artificial category system
based on the morphological similarity and historical function of each dog breed. Behavioural
comparisons at the breed-group level produced ambiguous results as to whether the his-
torical function still has an influence on the breed-typical behaviour. Recent genetic studies
have uncovered genetic relatedness between dog breeds, which can be independent from
their historical function and may offer an alternative explanation of behavioural differ-
ences among breeds. This exploratory study aimed to investigate the behaviour profiles
of 98 breeds, and the behavioural differences among conventional breed groups based on
historical utility and among genetic breed clusters. Owners of 5733 dogs (98 breeds) filled
out an online questionnaire in German. Breed trait scores on trainability, boldness, calm-
ness and dog sociability were calculated by averaging the scores of all individuals of the
breed. Breeds were ranked on the four traits and a cluster analysis was performed to explore
behavioural similarity between breeds.
We found that two of the behaviour traits (trainability and boldness) significantly dif-
fered both among the conventional and the genetic breed groups. Using the conventional
classification we revealed that Herding dogs were more trainable than Hounds, Working
dogs, Toy dogs and Non-sporting dogs; Sporting dogs were also more trainable than Non-
sporting dogs. In parallel, Terriers were bolder than Hounds and Herding dogs. Regarding
genetic relatedness, breeds with ancient Asian or African origin (Ancient breeds) were less
trainable than breeds in the Herding/sighthound cluster and the Hunting breeds. Breeds
in the Mastiff/terrier cluster were bolder than the Ancient breeds, the breeds in the Herd-
ing/sighthound cluster and the Hunting breeds. Six breed clusters were created on the
basis of behavioural similarity. All the conventional and genetic groups had representa-
tives in at least three of these clusters. Thus, the behavioural breed clusters showed poor
correspondence to both the functional and genetic categorisation, which may reflect the
effect of recent selective processes. Behavioural breed clusters can provide a more reliable
characterization of the breeds’ current typical behaviour.
© 2011 Elsevier B.V. All rights reserved.
Corresponding author. Tel.: +36 1 3812179; fax: +36 1 3812180.
E-mail addresses: (B. Turcsán), (E. Kubinyi), (Á. Miklósi).
1. Introduction
The dog (Canis familiaris) was the first domesticated
species, and descended from the gray wolf (Canis lupus)
at least 15,000 years ago (Savolainen et al., 2002; Dayan,
1994). Modern dog breeding over the past few hundred
years has generated great variation in morphology, physi-
0168-1591/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
62 B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70
ology and behaviour, which has giving rise to more than
400 dog breeds (Clutton-Brock, 1995) recognized today
by official dog kennel organisations around the world.
Due to modern breeding rules and strict breed standards,
dog breeds have become morphologically homogeneous,
genetically isolated breeding units (vonHoldt et al., 2010).
It is not clear whether the term “breed” refers only to the
genetic/morphological component or whether breeds are
also uniform in their behaviour. Breed differences have
been described predominantly in terms of behavioural
traits, which are recognized as derived features of context
independent behavioural functions, such as aggressive-
ness (Duffy et al., 2008) and nerve stability (Wilsson and
Sundgren, 1997a). However, researchers have also found
great individual variability within a single breed (e.g.
aggressiveness: Podberscek and Serpell, 1996 in English
Cocker Spaniel; nerve stability: Ruefenacht et al., 2002 in
German Shepherd Dog).
Surveys have been conducted to obtain behavioural
profiles of dog breeds by ranking the breeds on differ-
ent behaviour traits (Hart and Miller, 1985), or clustering
breeds on the basis of their behaviour (e.g. Bradshaw
and Goodwin, 1998; Hart and Hart, 1985; Takeuchi and
Mori, 2006). The concordance rate between two particular
surveys, which used almost identical methods for cluster-
ing breeds, was 50–60% (Bradshaw and Goodwin, 1998;
Takeuchi and Mori, 2006). This result could reflect a real
basis for breed-typical behaviours, but also suggests possi-
ble cultural differences. Thus our first aim is to characterize
dog breeds on the basis of their typical behaviour.
Behavioural differences between breeds are usually
explained by their historical function. Before the emer-
gence of dog kennel clubs at the end of the 19th century,
breeds (or certain type of dogs) were selectively bred to
optimize their performance in several tasks (e.g. herd-
ing, hunting, and guarding), which required selecting for
specific morphological and behavioural features. Represen-
tatives of a given breed are generally suitable for a specific
function (e.g. herding), due to breed specific behavioural
skills (see also Coppinger and Coppinger, 2001; Spady and
Ostrander, 2008). For example, livestock guarding dogs
should not show any predatory motor patterns (giving eye,
stalking and chasing) toward sheep or cows, while the pres-
ence of these behaviour patterns is important in herding
dogs (Coppinger and Schneider, 1995). Accordingly, breeds
with similar historical functions should behave generally
more comparably than breeds with different functions. One
problem which arises when using conventional grouping
methods to categorise breeds by function, is the lack of sci-
entific evidence on the history and function of most dog
breeds. The currently recognized breed groupings were
created by different national kennel club organisations,
like the Fédération Cynologique Internationale (FCI) or the
American Kennel Club (AKC), and are based on morpho-
logical similarity, anecdotal information about the breeds’
behavioural utility, and scarce historical evidence. How-
ever, dogs today are usually regarded as family members
or as companions in Western-cultures (Kubinyi et al., 2009;
Serpell, 2003) and are generally no longer utilised in their
original role. Behavioural traits, which had been the pri-
mary target for many hundreds of years, play little role
in the breed standards today (McGreevy and Nicholas,
1999). This may explain why the behavioural comparisons
at the breed-group level produced ambiguous results. For
example, Svartberg (2006) did not find behavioural differ-
ences in curiosity/fearlessness, aggression, playfulness, and
sociability among four FCI-based breed groups, whereas
Ley et al. (2009) reported differences in five questionnaire
scales in seven breed groups recognised by the Australian
National Kennel Council. Taken together, it is still an open
question whether the historical function of a breed affects
their behaviour. Thus, our second aim is to compare the
behaviour of breeds in conventionally recognized breed
Genetic relatedness could also account for behavioural
similarity among breeds (e.g. Takeuchi and Mori, 2006).
Many reports have provided evidence for genetic varia-
tion of behavioural traits in dog breeds (e.g. fearfulness:
Goddard and Beilharz, 1982, 1983; activity: Wilsson
and Sundgren, 1998). Accordingly, closely related breeds
should behave generally more similarly than genetically
more distant breeds. The application of modern genetic
methods based on similarity in DNA sequences has allowed
a more precise estimation of genetic relatedness among
breeds. However, this analysis has not revealed a true
phylogenetic relationship among breeds in the usual (evo-
lutionary) sense because of multiple cross-breeding events.
Nevertheless, recent genetic studies have been aimed at
analysing the hierarchical relationships between breeds.
For example, Parker et al. (2004) have generated four
genetic breed clusters from 85 breeds on the basis of 96
microsatellite loci. According to their analysis, a subset of
breeds with ancient Asian and African origins have split
off from the rest of the breeds with modern European ori-
gins, and shows the closest genetic relationship to the wolf.
The modern European breeds were later divided into three
clusters corresponding to the Mastiff, Herding and Hunting
breeds. A more recent study based on a larger sample size of
132 breeds (Parker et al., 2007) identified a fifth ‘Mountain’
cluster containing mostly large mountain dogs separated
from the Hunting cluster. More detailed analysis of breed
relationships revealed smaller closely related sub-clusters
within these five clusters, suggesting additional levels of
relatedness among some breeds (Parker et al., 2007). Lit-
tle is known about the effects of genetic relatedness on the
behaviour of dog breeds, thus, our third aim is to compare
the behaviour of breeds in different genetic breed clusters.
Traditionally two approaches have been used to char-
acterize the behaviour of dog breeds: breed rating (e.g.
Notari and Goodwin, 2007) and individual-based methods.
We followed the latter method, and used several individu-
als per breed and breeds per breed group. Individual-based
methods use two types of measurements: direct observa-
tional methods and questionnaire-based ratings of dogs
by their owners. Behavioural tests measure a restricted
set of objectively described behaviour units (e.g. growling
and tail wagging) in a few, controlled situations, whereas
questionnaire surveys are based on the owners’ knowl-
edge and familiarity with their dogs’ everyday behaviour.
The questionnaire method offers ease of data collection, a
larger and more diverse sample, and therefore meet the
requirements of this study. However there is some subjec-
B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70 63
tivity in these assessments, such as the differing experience
the owners have with dogs in general (Bennett and Rohlf,
2007, but see Tami and Gallagher, 2009). By combining the
responses of many independent owners, such individual
bias can be overcome (Jones and Gosling, 2005). The sec-
ond source of subjectivity is the possible influence of breed
stereotypes. Owners may tend to associate breeds or breed
groups with certain behaviour-types (i.e. assess their dogs’
behaviour on the basis of stereotypical beliefs). This sub-
jectivity seems to be unavoidable; however, as Kwan et al.
(2008) pointed out, the judgments of dogs do not simply
reflect breed stereotypes, but take into account each indi-
vidual’s behavioural features.
In this study, our main aims are (1) to characterize
dog breeds on four complex behavioural traits (trainabil-
ity, boldness, calmness and dog sociability) using owner
reported assessments on a large sample of dogs and (2)
to test whether dog breeds’ behavioural differences could
be ascribed to the breeds’ historical function (conventional
breed groups) and/or genetic relatedness. Since not much
research has been done on this topic and these issues we
addressed have not been clearly defined, this study should
be considered explorative by nature.
2. Materials and methods
2.1. Subjects
The analyses of the present study are based on a sub-
set of the database provided by Kubinyi et al. (2009) in
which 14,004 questionnaires were collected by the German
“Dogs” magazine (published by Living at Home Multi Media
GmbH, Hamburg, August 2007 issue) and the magazine’s
own website ( from August 2007
to January 2008. From this database we extracted all adult
pure-bred dogs whose breed presented at least 10 repre-
sentatives. Altogether, 5733 questionnaires from 98 breeds
were analysed. The dogs were on average 4.0 ±3.0 years
old, and 57.6% of them were males. Thirteen breeds were
represented by at least 100 individuals, and the most
frequent breed was the Labrador Retriever with 517 indi-
2.2. Procedure
We used an online questionnaire adapted for dogs
by Jesko Wilke based on a 48-item Human Personality
Inventory ( This question-
naire consists of 24 items in which owners are asked to
score their dogs using a 3-point scale. Previous results
(based on a larger sample from the original database) using
principal component analysis have revealed that 17 items
belonged to four components, labelled as trainability, bold-
ness, calmness and dog sociability (Kubinyi et al., 2009)
(Table 1). Dogs that scored low regarding the trainability
trait are described by their owners as uninventive and not
playful, whereas dogs that scored high on this trait are
regarded as intelligent and playful. Boldness was related
to fearful and aloof behaviour with a low score corre-
sponding to a high degree of fearfulness/aloofness, and vice
versa. The calmness trait describes the dogs’ behaviour in
stressful/ambiguous situations. A low score on this trait
indicated stressed and anxious behaviour in these situa-
tions, while a high score referred to calm and emotionally
stable dogs, according to the owner. Finally, dog sociability
refers to their behaviour toward conspecifics, with a low
score indicating a high tendency for bullying or fighting
and inversely high scores related to a low tendency. The
internal consistency and stability of these traits as well as
the test–retest reliability of the questionnaire was reported
earlier in Kubinyi et al. (2009).
Scores for individual traits were calculated by averag-
ing the scores from the variables representing each trait,
according to Kubinyi et al. (2009) (Table 1). The scores were
then averaged for each breed regarding each behavioural
trait in order to obtain the breed trait scores (Svartberg,
Hierarchical cluster analysis based on the breed trait
scores was used to investigate the behavioural similar-
ity between breeds. Hierarchical cluster analysis is an
exploratory method used to identify relatively homoge-
neous groups of cases based on selected characteristics. In
our sample, six breeds split off separately from the rest of
the breeds (see Appendix A) and the 92 remaining breeds
were divided into six clusters after visual examination of
the hierarchical structure.
The classification of breeds according to historical func-
tion was based on the internationally recognized system
of the American Kennel Club (AKC, Eight
breeds which are not recognized by the AKC (Bavarian
Mountain Hound, German Bracke, German Hunting Terrier,
Hovawart, Kromfohrländer, Landseer, Spanish Greyhound,
White Swiss Shepherd Dog), were assigned to whichever
AKC breed group most closely matched their classifica-
tion by the Fédération Cynologique Internationale (FCI, The 98 breeds present in this study were
therefore classified into seven groups (Table 2).
To analyse the effect of the breeds’ genetic relatedness
in some behaviour traits, we categorised the breeds into
five clusters according to Parker et al. (2007) (Table 3).
2.3. Statistical analyses
As the residuals of the breed behaviour trait scores
were not all normally distributed and the variances were
not homogeneous in all groups, nonparametric statistical
methods were used. Kruskal–Wallis tests with Dunn post-
hoc tests were used to compare the trait scores between
breeds, and between the seven conventional breed groups
and the five genetic breed clusters. A cluster analysis
based on the behavioural traits was performed using the
hierarchical agglomerative method. Distances between
breeds were calculated from the four traits by squared
Euclidean distance and breeds were clustered based on
the between-groups average linkage method. With this
method, a breed’s behaviour has to be within a certain level
of similarity to the cluster’s average to be included in that
cluster (Aldenderfer and Blashfield, 1984). Since the differ-
ent breed frequencies in our sample may bias the breed
trait scores, we randomly chose 10 individuals from each
breed and ran the analyses on this balanced sample as well.
SPSS 13.0 was used for all the analyses except the Dunn
64 B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70
Table 1
The 17 questions, belonging to four traits: trainability, boldness, calmness and dog sociability.
Trainability Calmness
Is ingenious, inventive when seeks hidden food or toy Is calm, even in ambiguous situations
Is intelligent, learns quickly Can be stressed easilya
Is very easy to warm up to a new toy Is emotionally balanced, not easy to rile
Often does not understand what was expected from him/her during playingaIs cool-headed even in stressful situations
Is not much interested except in eating and sleepingaIs sometimes anxious and uncertaina
Boldness Dog sociability
Is rather cool, reservedaFights with conspecifics frequentlya
Is unassertive, aloof when unfamiliar persons enter the homeaIs ready to share toys with conspecifics
Is bullying with conspecificsa
Is sometimes fearful, awkwardaGets on well with conspecifics
aScoring was reversed.
Table 2
Breed distribution according to conventional categorisation.
Conventional group name and description Number of breeds Number of individuals
Sporting dogs Include pointers, retrievers, setters and spaniels; mostly
used for cooperative hunting.
15 1197
Hounds Include scenthounds, greyhounds and dachshunds; being
used for independent hunting.
11 528
Working dogs Were bred to perform such jobs as guarding livestock or
pulling sleds.
20 1025
Terriers Middle or small sized breeds, used for independent
14 808
Toy dogs Small sized breeds with the main function:
11 561
Non-sporting dogs Diverse group in terms of size and utility 12 492
Herding dogs Middle or large sized breeds used for control the
movement of other animals.
15 1122
All groups 98 5733
post-hoc tests, for this we used GraphPad Instat statistical
3. Results
3.1. Breed differences in behavioural traits
There were significant differences between breeds
in all four traits (Kruskal–Wallis test, N= 5733, d.f. = 97,
trainability 2= 641.405; boldness 2= 417.126; calmness
2= 455.005; dog sociability 2= 687.035, P< 0.001 for all).
The breeds were ranked on the basis of these traits
(Appendix A). The five most popular breeds (N200: Bea-
gle, German Shepherd Dog, Golden Retriever, Jack Russell
Terrier, and Labrador Retriever) had no higher or lower
scores than the population mean ±SD in calmness and
trainability. However, the Beagle and Labrador Retriever
scored higher than the mean on dog sociability, while the
German Shepherd Dog and Jack Russell Terrier obtained
lower scores. Jack Russell Terriers and Labrador Retrievers
also scored higher on boldness.
3.2. Behavioural similarity between breeds
Breeds were clustered on the basis of their behaviour
using hierarchical cluster analysis. Six breeds (three breed
pairs) with extreme trait scores split off from the other
breeds (Appendix A). The first pair consisted of the
Newfoundland and the Landseer, known to be strongly
genetically related. The AKC and the Kennel Club in the
Table 3
Breed distribution according to the genetic relatedness (Parker et al., 2007).
Genetic cluster name and description Number of breeds Number of individuals
Ancient breeds Breeds with ancient Asian or African
origin, mainly primitive type dogs
7 192
Mastiff/terrier cluster Mastiff-type breeds or breeds with
mastiff-type ancestors and terriers
13 1019
Herding/sighthound cluster Breeds used as herding dogs and
10 674
Hunting breeds Breeds with relative recent European
origin, primarily different hunting dogs
29 1974
Mountain cluster Large mountain dogs and a subset of
11 958
All groups 70 4817
B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70 65
UK classify them as coat colour varieties of a single breed,
although the FCI recognizes them as distinct breeds. Both
breeds were highly ranked in boldness, calmness and dog
sociability. The next pair comprised of the Akita and the
German Bracke and both breeds scored extremely low in
regards to the dog sociability trait. The third pair, the Ger-
man Pincher and the Spanish Greyhound, only slightly
related to each other. Both breeds scored low on calmness,
and the Spanish Greyhound ranked also low on boldness
and trainability and high on dog sociability.
The remaining 92 breeds were divided into six clus-
ters with 2–32 breeds in each cluster (Table 4), according
to the dendrogram. The clusters differed from each other
in each of the four traits (Kruskal–Wallis test, N= 92,
d.f. = 5, trainability 2= 43.409; boldness 2= 53.208, calm-
ness 2= 45.790, dog sociability 2= 49.507, P< 0.001, for
all) and were characterized as low, medium and high on
each trait, based on the post-hoc differences between them
(Table 4).
3.3. Differences among conventional breed groups
Significant differences in the trainability and bold-
ness scores were observed between the breed groups
(Kruskal–Wallis test, N= 98, d.f. = 6, trainability 2= 31.025,
P< 0.001; boldness 2= 19.325, P= 0.004); nevertheless,
the differences between the groups regarding calmness and
dog sociability traits were not significant (Kruskal–Wallis
test, N= 98, d.f. = 6, calmness 2= 11.522, P= 0.074; dog
sociability 2= 12.111, P= 0.06) (Fig. 1A and B).
According to the post-hoc tests, Herding dogs were
reported by their owner to be more trainable than Hounds
(P< 0.01), Working dogs (P< 0.01), Toy dogs (P< 0.05) and
Non-sporting dogs (P< 0.001). Sporting dogs were also
more trainable than Non-sporting dogs (P< 0.05).
Terriers scored higher on boldness than Hounds
(P< 0.01) and Herding dogs (P<0.05) (Fig. 1A).
3.4. Genetic relatedness
The five genetic clusters (identified by Parker
et al., 2007) differed also in trainability and bold-
ness (Kruskal–Wallis test, N= 70, d.f. = 4, trainability
2= 10.153, P= 0.038; boldness 2= 14.497, P= 0.006).
The cluster of Ancient breeds was less trainable than the
Herding/sighthound cluster and the cluster of Hunting
breeds (P< 0.05 for both).
The Mastiff/terrier cluster was bolder than the Ancient
breeds, the Herding/sighthound cluster and also the Hunt-
ing breeds (P< 0.05 for all) (Fig. 2A).
No significant differences in calmness and dog
sociability traits were found between these clusters
(Kruskal–Wallis test, N= 70, d.f. = 4, calmness 2= 7.915,
P= 0.095; dog sociability 2=2.785, P= 0.594) (Fig. 2B).
The analyses of the random sample of 980 individu-
als (10 individuals per breed, as mentioned in Section 2)
led to the same differences between genetic breed clusters
and similar differences between the conventional breed
groups. The differences in calmness and dog sociability
traits among conventional breed groups were significant in
this sample compared to the trend difference observed in
1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7
Sporting dogs
Herding dogs
Toy dogs
Working dogs
1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7
Dog Sociability
Working dogs
Sporting dogs
Non-sporting dogs
Toy dogs
Fig. 1. Behaviour profiles of the conventional breed groups in (A) train-
ability and boldness and (B) calmness and dog sociability traits. Data
points are group averages, the diameters of the ellipses represent the SE
of trainability/calmness, the heights the SE of boldness/dog sociability.
Dashed lines represent the population means. Kruskal–Wallis test showed
significant differences between the groups in trainability and boldness
and trend differences in calmness and dog sociability traits.
the whole sample. Nevertheless, this finding supports that
the different breed frequencies in this study only minimally
influenced the results.
4. Discussion
The main focus of this study was to discover the typ-
ical behaviour of dog breeds and specific breed groups.
Our aims were (1) to characterize a large number of
breeds and explore their behavioural similarity and diver-
gence, and (2) to test whether dog breeds’ behavioural
differences could be ascribed to the breeds’ conventional
66 B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70
Table 4
Clusters of breeds from the cluster analysis based on the trainability, boldness, calmness and dog sociability traits. The numbers links to the breeds represent
the conventional breed groups and genetic clusters, respectively. The characteristic features of each cluster are presented.
Cluster 1 high calm, medium trainable, high sociable, high bold
4;2 Airedale Terrier 3;5 Greater Swiss Mountain Dog 3;5 Saint Bernard
1;4 American Cocker Spaniel 5;. Havanese 7;3 Shetland Sheepdog
2;4 Beagle 1;4 Irish Setter 5;4 Shih Tzu
7;3 Bearded Collie 1;2 Labrador Retriever 3;1 Siberian Husky
3;5 Bernese Mountain Dog 3;5 Leonberger 1;. Small Munsterlander
6;2 Bulldog 4;4 Miniature Schnauzer 4;2 Soft Coated Wh. Terrier
5;4 Cavalier King Charles Spaniel 7;3 Old English Sheepdog 4;2 Staffordshire Bull Terrier
6;2 French Bulldog 5;4 Pekingese 2;3 Whippet
1;4 Golden Retriever 1;4 Pointer
1;4 Gordon Setter 5;4 Pug
Cluster 2 low calm, high trainable, low sociable, low bold
4;2 American Staffordshire Terrier 1;5 English Cocker Spaniel 4;. Parson Russell Terrier
7;3 Australian Shepherd 7;. Entlebucher Mountain Dog 6;5 Poodle
2;. Bavarian Mountain Hound 4;. German Hunting Terrier 3;5 Rottweiler
7;3 Belgian Malinois 7;5 German Shepherd Dog 3;4 Standard Schnauzer
7;3 Border Collie 1;. German Wirehaired Pointer 6;1 Tibetan Terrier
4;2 Border Terrier 3;4 Giant Schnauzer 1;4 Vizsla
3;2 Boxer 3;3 Great Dane 4;2 Welsh Terrier
4;4 Cairn Terrier 3;. Hovawart 4;4 West Highland Wh.Terrier
2;4 Dachshund 4;2 Irish Terrier 2;. Wirehaired Dachshund
6;4 Dalmatian 4;4 Jack Russell Terrier 6;4 Wolfspitz
3;4 Doberman Pinscher 5;5 Miniature Pinscher
Cluster 3 high calm, high trainable, high sociable, high bold
1;4 Flat-Coated Retriever 1;4 German Shorthaired Pointer
Cluster 4 low calm, high trainable, medium sociable, low bold
7;. Appenzeller Sennenhund 5;. Kromfohrländer 2;5 Rhodesian Ridgeback
7;. Beauceron 2;. Miniature Dachshund 6;1 Shiba Inu
7;. Briard 5;4 Miniature Poodle 1;4 Weimaraner
7;3 Collie 7;. Polish Lowland Sheepdog 7;. White Swiss Shepherd
2;4 Ibizan Hound 7;. Pyrenean Shepherd
Cluster 5 low calm, low trainable, low sociable, medium bold
1;4 Brittany 6;. German Spitz 5;. Yorkshire Terrier
4;. Bull Terrier 5;5 Maltese
5;4 Chihuahua 3;2 Perro de Presa Canario
Cluster 6 high calm, low trainable, medium sociable, low bold
3;1 Alaskan Malamute 6;. Coton de Tulear 6;. Eurasier
3;. Anatolian Shepherd Dog 3;. Dogue de Bordeaux 2;3 Irish Wolfhound
6;1 Chinese Shar-Pei 1;4 English Setter 6;1 Lhasa Apso
grouping based on historical function and/or genetic relat-
edness. We derived complex breed-related behavioural
traits by averaging the behaviour scores of individual dogs
within a given breed. This was based on the assumption
that the complex behavioural traits might distinguish not
only individual dogs, but also typical for larger population
of dogs representing breeds or special breed groupings.
Individual dog behaviour was measured by an owner-
reported assessment, and four behavioural traits were
obtained, namely trainability, boldness, calmness and dog
sociability. Similar traits were previously detected by sev-
eral other researchers (e.g. trainability: Bradshaw and
Goodwin, 1998; Hsu and Serpell, 2003; boldness: Svartberg
and Forkman, 2002; neuroticism (reverse to our calmness):
Ley et al., 2009; dog-directed aggression (reverse to our dog
sociability): Hsu and Serpell, 2003). The four behavioural
traits found in this study correspond to the seven person-
ality dimensions of dogs summarized in Jones and Gosling
(2005) (see also in Kubinyi et al., 2009).
4.1. Breed differences in behavioural traits
Significant breed differences were observed in all the
four traits. The behavioural profiles of the breeds are to
some extent in accordance with the reports of other stud-
ies, but there is some dissimilarity. For example in the
study of Duffy et al. (2008) the Akita, Dachshund, Chi-
huahua and Jack Russell Terrier were reported to be highly
aggressive toward other dogs, whereas the Whippet, Col-
lie and Bernese Mountain Dog were at the opposite end of
the scale. Accordingly, the former four breeds ranked low
in dog sociability in this study (lower than the population
B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70 67
Fig. 2. Behaviour profiles of the genetic breed clusters in (A) trainability
and boldness and (B) calmness and dog sociability traits. Data points are
group averages, the diameters of the ellipses represent the SE of trainabil-
ity/calmness, the heights the SE of boldness/dog sociability. Dashed lines
represent the overall mean of the factors. Kruskal–Wallis test showed sig-
nificant differences between the groups in trainability and boldness and
trend difference in calmness traits.
mean ±SD), but from the later three, only the Whippet and
Bernese Mountain Dog were ranked high, and the Collie
was in the middle section of the dog sociability rank order.
Breed standards of kennel clubs usually contain a short
description about the favourable behavioural character-
istics of a given breed. For example, according to its FCI
breed standard, Anatolian Shepherds are “steady and bold
without aggression, naturally independent, very intelligent
and tractable, proud and confident” ( More-
over, one of the eliminating faults of this breed is signs
of aggressiveness or shyness. Contrary to this description,
the Anatolian Shepherd breed in this study was described
as low trainable, highly shy and not sociable with other
dogs. Another example is the Spanish Greyhound (Galgo
Espanol), which, according to its breed standard, should not
be aggressive or overly shy (one of the eliminating faults),
here was found to be the least bold breed of all. These exam-
ples show that even the vague characteristics described in
the breed standards do not always reflect reality.
The dendrogram of the cluster analysis based on the
four behavioural traits (Appendix A) illustrates the highly
complex relationship between breeds. We divided the
breeds into six clusters which either show or do not show
strong correspondence to the conventional categories or
the genetic breed clusters of Parker et al. (2007). There
is some correspondence with the behavioural breed clus-
ters in other studies. For example, eight of the nine breeds
which were represented in our sample from the Svartberg’s
(2006) Cluster 1 also clustered together in this study. The
Border Collie and Australian Shepherd had a highly sim-
ilar behavioural profile which corresponded well to their
genetic and functional similarity. In contrast, the Shetland
Sheepdog and the Collie were related both in their his-
torical function and genetically (Neff et al., 2004; Parker
et al., 2007) but were clustered far from each other in their
behavioural profiles. Given the explorative nature of our
study and the applied statistical method, the behavioural
relationship between breeds described in this study should
be interpreted with caution; more studies are needed to
confirm our findings.
4.2. Conventional categorisation
Dog breeds were grouped based on the systems pro-
vided by two internationally recognized kennel clubs,
presupposing that these systems reflect the breeds’ histor-
ical function. The seven groups differed in both trainability
and boldness traits.
Trainability: Herding dogs were more trainable than
Hounds, Working dogs, Toy dogs and Non-sporting dogs.
Sporting dogs were also more trainable than Non-sporting
dogs. Similar behavioural differences were previously
shown in several surveys. Indeed, both Seksel et al. (1999)
and Ley et al. (2009) found that breeds from the Pointing
dogs group (here classified as Sporting dogs) were highly
trainable, while Toy dogs scored low on that scale. Addi-
tionally, Ley et al. (2009) reported that Hounds were also
relatively less trainable compared to Pointing and Herd-
ing dogs. Serpell and Hsu (2005) constructed a rank order
among different breeds and found that the most trainable
breeds were either the representatives of Herding or Point-
ing groups. The authors explained their results on the basis
of the cooperative versus independent type of work that
the breeds were originally bred for. Indeed, Herding and
pointing dogs were bred for cooperative tasks with con-
tinuous visual contact with their human partner, while
hounds were to hunt independently, out of the view of
humans (see Gácsi et al., 2009). The Working dog group
consists of sled dogs, guarding dogs and livestock guard-
ing dogs, which require some human guidance to execute
their tasks successfully, however most of these breeds were
selected for independent working.
68 B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70
Other factors, such as the differences regarding the
general size or physical abilities of dogs could also be
responsible for the breed/breed-group differences (Helton,
2010). For instance, the low trainability of the Toy dogs
group could be explained by the typically small physical
size of this group (Bennett and Rohlf, 2007), since larger
dogs are more likely to attend formal obedience training,
possibly because disobedience or behavioural problems
could be more serious in larger dogs (Kobelt et al., 2003).
Non-sporting dogs appear to be the most difficult to
train. This group contains highly diverse breeds in terms
of size or historical function and is dominated by breeds
with ancient Asian origin. The close genetic relatedness of
these breeds with wolves (Parker et al., 2004) may account
partly for the low trainability skills observed in this group.
Boldness: Terrier breeds scored higher on boldness than
Hounds and Herding dogs, in line with previous stud-
ies in which terriers are described as typically energetic,
excitable and reactive dogs (Hart, 1995; Ley et al., 2009;
Scott and Fuller, 1965).
The historical function and utility of dog breeds seem
to still have an effect on breeds’ typical behaviour. How-
ever, explaining breed differences in terms of their function
solely would presuppose that dogs are bred on the basis
of their performance regarding these functions; which is
usually not the case. Indeed, in modern dog breeding, most
animals are selected on the basis of their success and judge-
ment at dog shows. Dogs are evaluated at these shows
according to strict breed standards which reflect mostly
on morphological requirements. It is however important
to note that breeds are often separated into working
and show “lines” or “types”, based on divergent selec-
tion criteria (e.g. Border Collie, Chang et al., 2009). In
working lines, the performance plays the most important
role for breeding. Accordingly, the behaviour of working
line individuals could differ from that of the show lines
(Duffy et al., 2008).
The differences between the conventional breed groups
in calmness and dog sociability were rather small (trend
differences in the whole sample and significant group dif-
ferences in the random sample), although there were large
differences between breeds in both traits. These charac-
teristics are not strictly related to the breeds’ original
function, however, they might have been important for
some breeds. For example, some breeds from the Hound
group which were used for hunting in groups should show
low aggression toward conspecifics as a requirement in
these breeds; whereas other functions like herding might
not have required direct selection for tolerant behaviour
toward conspecifics.
However, the differences between the conventional
breed groups in trainability and boldness are in con-
trast with the findings of Svartberg (2006), who did not
find any breed group differences in four traits measured
by behavioural tests. He suggested that recent selection
criteria are more important in shaping the behaviour
of dog breeds than their historical function. The differ-
ences between the two studies might reflect real breed
differences between countries due to genetic isolation
(Notari and Goodwin, 2007) or the different attitudes
toward dog breed standards and present utility as well
as various environmental effects (e.g. keeping condi-
tions, neutering) in each country (Houpt et al., 2007).
Different breed representations between countries and
breed popularity could also potentially affect dog breeds’
behaviour (Svartberg, 2006). However, it is also possi-
ble that our results only reflect the owners’ stereotypic
beliefs about breeds and breed groups of different historical
4.3. Genetic relatedness
Breeds were grouped into five clusters (Parker et al.,
2007) presupposing that the classification based on 96
microsatellite loci reliably mirrored the breeds’ genetic
relatedness. To our knowledge the present analysis is
the first to investigate the effect of genetically supported
grouping on the behavioural traits of dogs. The five clusters
differed in trainability and boldness traits.
Ancient breeds were less trainable than breeds in the
Herding/sighthound cluster and Hunting breeds. Breeds
from the Mastiff/terrier cluster were bolder than the
Ancient breeds, the breeds from the Herding/sighthound
cluster and the Hunting breeds.
It seems that the genetic separation of the Ancient
breeds from the other four breed clusters representing
modern European breeds is paralleled by differences in
behaviour. The Ancient breeds show the closest genetic
relationship with the wolf (Parker et al., 2007) which might
explain some part of these differences. The high score
in the boldness trait of the Mastiff/terrier genetic cluster
parallels the high boldness score of the Terriers and Work-
ing dogs in the conventional categories (Fig. 1). There is
some evidence in the literature on the genetic basis of
behavioural traits which are related to fearfulness (e.g.
Murphree et al., 1969; Peters et al., 1967), for example,
Goddard and Beilharz (1983) found relatively strong (0.46)
heritability for this trait. However, much lower heritabil-
ity was reported for ‘courage’ in German Shepherd and
Labrador Retriever (0.25 and 0.28, respectively) (Wilsson
and Sundgren, 1997b), and for boldness in German Shep-
herd and Rottweiler (0.25–0.27, respectively) (Strandberg
et al., 2005; Saetre et al., 2006). Finally, trainability showed
even lower heritability (0.01–0.16, reviewed in Ruefenacht
et al., 2002).
The differences found between the genetic clusters were
not independent from those found between the conven-
tional breed groups, since genetic relatedness is often
associated with morphological and functional similarity
and shared geographic origin (Parker et al., 2004; vonHoldt
et al., 2010). For example, conventional categorisation
revealed that Terriers were bolder than Herding dogs. In
parallel dogs in the genetic cluster of Mastiff/terrier breeds
scored higher than the breeds in the Herding/sighthound
cluster. It is not evident yet, if there is a causal relationship
between these two results and if so, what is the direction
of the causality. Overlaps between the conventional and
the genetic groups are not obvious, because more recent
breeds, especially in the Working dogs and Toy dogs groups
were frequently bred from combinations of other breeds,
independently of their historical function and genetic relat-
edness (vonHoldt et al., 2010).
B. Turcsán et al. / Applied Animal Behaviour Science 132 (2011) 61–70 69
One limitation of the present study was that the own-
ers’ assessment of their dog could be biased by subjectivity
and stereotypical beliefs. Another limitation is that dog
keeping practices, which could affect the dogs’ behaviour
(Bennett and Rohlf, 2007; Kobelt et al., 2003), may vary sys-
tematically among breeds, producing false similarities or
discrepancies in the perception of their typical behaviour.
However, the diversity and the large number of dogs inves-
tigated in this study may help to minimize these biases.
5. Conclusion
We found large differences among dog breeds in four
behavioural traits. Our results showed that the differences
in breed-specific behaviour in trainability and boldness are
partly determined by genetic factors and differences in
the historical function of the breeds. However, breed clus-
ters with similar behavioural characteristics corresponded
neither to the presently recognized functional (conven-
tional) classification nor to the genetic clusters of breeds.
Behavioural divergence of seemingly related dog breeds
could be associated to either cross-breeding with other
breeds characterised by different behavioural traits (e.g.
selection for toy dogs, vonHoldt et al., 2010), ceasing selec-
tion for the original function, or changes in function that
was associated with a novel selective environment. Other
factors like early period of socialization, dog keeping prac-
tices, and the behaviour of the owner might also play an
important role in shaping the dogs behaviour, and these
factors can modify the behaviour of individual dogs from
the core characteristics typical of their breeds. Never-
theless, our study, although explorative by nature, might
help owners to choose the appropriate breed as a pet on
the basis of real breed-typical behaviour, which is bene-
ficial not only for the owner but also for the welfare of
the dog. Studies such as this one may be significant in
contrasting the typical behaviour of “real” dog breed pop-
ulations with the often ‘ad hoc’ description used by kennel
clubs. It would be advantageous to use such data to make
more precise official breed descriptions to help ensure
a better match between a prospective owner and their
The authors are grateful to Jesko Wilke, journalist
associated with the German DOGS Magazine, for his coop-
eration in developing the questionnaire and for making this
study possible; and Krisztina Nagy from the Szent István
University for her help in analyzing the data. We also thank
Lisa Wallis for correcting the English in the manuscript.
The research was funded by the FP7-658 ICT-2007 (LIREC-
215554), the Hungarian Scientific Research Fund (K 84036)
and the Bolyai Foundation of the Hungarian Academy of
Appendix A. Supplementary data
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  • ... Early experience has been found to have a long-term effect on the personality of dogs (27)(28)(29). Additionally, several studies have established differences in personality between individuals belonging to dog breeds or breed groups (30)(31)(32), as well as between the typical personality of pure breed and mixed breed dogs (33). ...
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    The dog has been suggested as a possible model for personality development over the lifespan, however, we know little about how aging may shape their personality or the magnitude of age-related changes. Previously we established that aging influences multiple dog demographics, which could also affect how personality traits change across different age periods. A demographic questionnaire and the Dog Personality Questionnaire were completed for a cross-sectional sample of 1,207 adult dogs living in Hungary (M age = 7.71, SD = 4.12), split into six different age groups. Results revealed three of the five factors showed significant age effects. Activity/Excitability decreased with age, and whilst Responsiveness to training also decreased, only dogs older than 12 years differed significantly from the other groups. Aggressiveness toward animals showed a quadratic trajectory peaking in dogs aged 6-10 years. The greatest magnitude of age-related change was detected between late senior and geriatric ages, likely caused by compensatory behavioral changes to biological aging and owner attitudes to aging. When the models were rerun including the other explanatory variables, age group was no longer significant for the Responsiveness to training trait. The amount of time spent interacting/playing with the owner partially mediated the relationship between age and this trait, implying that interventions to increase play and training motivation may alleviate the negative effects of aging on dogs' trainability. Fifteen out of 28 explanatory variables were significantly associated with at least one of the five factors [weight, breed (pure/mixed breed), sex, off-leash activity, diet, previous trauma, age of dog when arrived in the household, play, dog training activities, number of known commands and dog obedience tasks]. Similarly to humans, dogs that had previously experienced trauma scored higher in fearfulness and aggression. A higher level of basic obedience was linked to some desirable dog personality traits (lower Fearfulness and Aggression, and higher Activity/Excitability and Responsiveness to training). Regardless of the direction of this relationship, obedience is an important aspect contributing to dog personality questionnaires and the dog-owner relationship. This study is unique in that it considered a wide variety of demographic variables which are influenced by aging.
  • ... Las 400 razas existentes presentan diferencias en su morfología y comportamiento, con una relación genética independiente de su función histórica (Turcsán, Kubinyi & Miklósi, 2011). La evolución de las razas preexistentes los clasificó según su fin zootécnico en "perros de servicio", desempeñándose como perros guía de invidentes, de asistencia, detectores de sustancias, búsqueda y rescate, entre otros (Hare & Tomasello, 2005). ...
  • ... Bold dogs (compared to shy dogs) had better resistance to diseases in a dog shelter, which is a stressful and very infectious environment. 26 As boldness and sociability differ between different breeds, [27][28][29] could it also be interpreted that some breeds are more resilient in facing adversities compared to others? The Labrador is among the boldest and most social (extraversion) dog breeds studied 12 and is also the most popular dog breed as a family pet in several countries, as well as highly used also as a working dog. ...
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    Katriina Tiira1,2 1SmartDOG, Riihimäki 11130, Finland; 2Department of Equine and Small Animal Medicine, University of Helsinki, Helsinki, FinlandCorrespondence: Katriina TiiraDepartment of Equine and Small Animal Medicine, University of Helsinki, PO Box 57, Helsinki FI-00014, FinlandEmail Katriina.tiira@helsinki.fiAbstract: What are the key factors of psychological resilience in dogs? Why do some individuals recover swiftly from neglect, abuse or several years of harsh kennel environments, while some seem to be permanently traumatized by much milder adverse experiences? Resilience is a concept seldom discussed in canine studies; however, many studies have identified risk factors (both environmental and genetic) for developing anxieties, aggression or other behavioral problems. These studies also indicate several factors that may act as protective agents against life adversities. In this paper, I will present some of the most commonly identified key factors of resilience in other species and discuss what has been found in dogs. This paper is an attempt to raise focus on the positive key factors in a dog’s life that are important for dog welfare, a healthy psychological outcome and are also important building blocks of a happy and well-behaving pet.Keywords: resilience, dog, stress
  • ... Few studies have applied a control based on a phylogenetic tree of dogs that could be estimated from genetic similarity (Overall, Tiira, Broach, & Bryant, 2014;Wayne & Ostrander, 2007), but the meaning of phylogeny in dogs has been suggested to be dubious (e.g. Careau, Reale, Humphries, & Thomas, 2010;Geiger et al., 2017;Turcsán et al., 2011). ...
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    Among‐population variance of phenotypic traits is of high relevance for understanding evolutionary mechanisms that operate in relatively short timescales, but various sources of non‐independence, such as common ancestry and gene flow can hamper the interpretations. In this comparative analysis of 138 dog breeds, we demonstrate how such confounders can independently shape the evolution of a behavioral trait (human‐directed play behavior from the Dog Mentality Assessment project). We combined information on genetic relatedness and haplotype sharing to reflect common ancestry and gene flow, respectively, and entered these into a phylogenetic mixed model to partition the among‐breed variance of human‐directed play behavior while also accounting for within‐breed variance. We found that 75% of the among‐breed variance was explained by overall genetic relatedness among breeds, while 15% could be attributed to haplotype sharing that arises from gene flow. Therefore, most of the differences in human‐directed play behavior among breeds have likely been caused by constraints of common ancestry as a likely consequence of past selection regimes. On the other hand, gene flow caused by crosses among breeds has played a minor, but not negligible role. Our study serves as an example of an analytical approach that can be applied to comparative situations where the effects of shared origin and gene flow require quantification and appropriate statistical control in a within‐species/among‐population framework. Altogether, our results suggest that the evolutionary history of dog breeds have left remarkable signatures on the among‐breed variation of a behavioral phenotype.
  • ... The dogs were selected because of their impressive olfactory system and because they are also relatively small, calm, highly trainable, and highly sociable. 12 One dog, Canine ...
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    Context: Early detection provides the best opportunity for lung cancer survival; however, lung cancer is difficult to detect early because symptoms do not often appear until later stages. Current screening methods such as x-ray and computed tomographic imaging lack the sensitivity and specificity needed for effective early diagnosis. Dogs have highly developed olfactory systems and may be able to detect cancer in its primary stages. Their scent detection could be used to identify biomarkers associated with various types of lung cancer. Objective: To determine the accuracy of trained beagles' ability to use their olfactory system to differentiate the odor of the blood serum of patients with lung cancer from the blood serum of healthy controls. Methods: Over the course of 8 weeks, operant conditioning via clicker training was used to train dogs to use their olfactory system to distinguish blood serum from patients with malignant lung cancer from blood serum from healthy controls in a double-blind study. After training, non-small cell lung cancer and healthy control blood serum samples were presented to the dogs, and the sensitivity and specificity of each dog were analyzed. Results: Four dogs were trained for the study, but 1 was unmotivated by training and removed from the study. Three dogs were able to correctly identify the cancer samples with a sensitivity of 96.7%, specificity of 97.5%, positive predictive value of 90.6%, and negative predictive value of 99.2%. Conclusion: Trained dogs were able to identify non-small cell lung cancer samples from healthy controls. The findings of this study provide a starting point for a larger-scale research project designed to explore the use of canine scent detection as a tool for cancer biomarkers.
  • ... In addition, dogs' performance in a cognitive task may not depend merely on mental abilities but also on factors such as temperament and personality. Turcsán et al. (2011) showed how the historical function and utility of dog breeds seem to still have an effect on breed typical behaviour: Mastiff-type breeds gained highest score in boldness traits compared to herding/hunting breeds. This trait of personality could partially explain their perseverance in the choice of the large quantity of food in spite of the experimenter's suggestions. ...
    The aim of this work was to investigate the effect of dog breed groups, i.e., primitive, hunting/herding and Mastiff like (Study 1) and development, i.e., 4-month-old puppies vs adults (Study 2) on a quantity discrimination task. The task consisted of three conditions: C1—dogs were asked to choose between a large and a small amount of food; C2—the same choice was presented and dogs could choose after having witnessed the experimenter favouring the small quantity. C3—similar to C2 but the plates had two equally small food quantities. Study 1 revealed that dogs in the hunting/herding group were significantly more likely than Mastiff-like group to choose the small quantity indicated by the person over the large one, although all dog groups chose the large quantity over the small when they had a free choice. These results are consistent with the hypothesis that hunting/herding breeds have been selected for working in cooperation with humans and thus may be more sensitive to human social communicative cues than other breeds. In Study 2, results showed that 4-month-old puppies performed at chance level in C1, whereas in C2 both adults and puppies conformed to the experimenter’s choice. In C3, adults followed the experimenter significantly more than puppies, although puppies still followed the experimenter above chance. Overall, domestic dogs seem to rely heavily on social communicative cues from humans, even when the information contradicts their own perception. This tendency to respond to human social cues is present, although at a lesser extent already at 4 months.
  • Article
    This paper reviews the history of the establishment of dog breeds, summarizes current health and resultant welfare problems and makes some positive suggestions for their resolution. Some breed standards and selection practices run counter to the welfare interests of dogs, to the extent that some breeds are characterized by traits that may be difficult to defend on welfare grounds. Meanwhile, little selection pressure seems to be exerted on some traits that would improve animal welfare and produce dogs better suited to modern society. Unfortunately, the incidence of certain inherited defects in some breeds is unacceptably high, while the number of registered animals of certain breeds within some countries is so low as to make it almost impossible for breeders to avoid mating close relatives. There are several constructive ways to overcome these challenges. Breed associations can ensure that reduction of welfare problems is one of their major aims; they can review breed standards; they can embrace modern technology for animal identification and pedigree checking; they can allow the introduction of 'new' genetic material into closed stud-books; and they can encourage collaboration with geneticists in identifying and using DNA markers for the control of inherited disorders. There should be a concerted effort to produce and evaluate as companion animals first-cross (F1) hybrids from matings between various pairs of breeds. Finally, geneticists must learn to communicate their science better and in a language that non-geneticists can understand.
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    Many dogs are found to be unsuitable for training as guide dogs for the blind. Consequently the Royal Guide Dogs for the Blind Association of Australia has embarked on a breeding program to produce a strain of labrador dogs which is suitable for guide dog training. The most common reasons for rejecting dogs are fearfulness, dog distraction, excitability, health and physical reasons and hip dysplasia. The selection program seems to have been successful in improving the success rate mainly by lowering fearfulness, but there has not been a continuing improvement. This is probably due to continual introduction of dogs from other populations into the breeding program. Males suffer from a higher rejection rate due to dog distraction and a lower rejection rate due to fearfulness and excitability than females, so that there is little sex difference in overall success rate. The heritability of success (0.44) is high enough to predict further progress from selection, again mainly against fearfulness. Variation in environment prior to 6 weeks of age, in age when dogs were placed into a private home and in age when males were castrated, had little effect on the success rate.
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    Domestic dogs in the Near East have been identified on the basis of cultural criteria, but identifications based on morphological criteria are controversial. Measurements of carnassial teeth and of the facial region of the cranium and mandible reveal that wolf/dog remains from the Natufian (c. 12,000 BP) and later cultures of Israel exhibit a morphological pattern that is the opposite of that expected under natural selection, but that conforms well to that expected in early domestication. The Geometric Kebaran wolves preceding the Natufian domesticates are very large individuals, probably in response to the climatic conditions of the period, and this may indicate one of the following: (a) the wolves domesticated in Israel were of a large race, contrary to previous theories on the roots of dog domestication; (b) the dog was domesticated at a period earlier than the Natufian.
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    The aims of this paper are first to review scientific and ethical considerations regarding the welfare of dogs in the US, UK, Czech Republic, Spain and Japan, as well as to examine research perspectives of people working within the field of applied ethology. As a guide, the Five Freedoms should be considered for dogs as with other species of animals [Brambell, F.W.R., 1965. Brambell, Report on the Technical Committee to Enquire into the Welfare of Livestock Kept Under Intensive Husbandry Systems, HMSO, London]. With pet dogs the use of drugs or shock therapy, neutering, caging, debarking and euthanasia for behaviour problems are all controversial not only among the general public but also among specialists in behavioural therapy. Breeding systems, evaluation of aptitudes, socialisation, training methods, and retired dogs’ lives are the major welfare concerns related to working dogs. Even within European countries, e.g. Spain, the Czech Republic, and the UK, there are different standards and attitudes towards dogs in each country. This suggests that scientific evidence and cultural consideration may be necessary to improving the welfare of companion animals. Different situations which exist between countries should be considered when seeking to establish international standards, however, these may also provide evidence for causes and inform possible modifications regarding canine behavioural issues. Sections contributed by authors from five countries discuss current issues and aim to produce a better understanding of local behaviour and welfare factors. It is concluded that there are still many welfare issues surrounding companion and working dogs internationally and further collaborative investigation is required in seeking to improve these.
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    The selection of dog breeds for functionality has progressively lost its importance and behavioural attributes originally selected are often now considered problems in household contexts. This study investigated behavioural characteristics of the 49 most popular breeds of dogs in Italy. Questionnaires were sent to 112 Italian veterinarians and 56 non-veterinarians (trainers, behaviour counsellors and animal charity officers) who rated breed behavioural characteristics and compared the behaviour of males and females. Females were considered more trainable for obedience, more demanding of affection and more housetrainable. Males were rated higher than females for all other traits except playfulness and general activity. Principal factor analysis with varimax rotation generated two principal factors (labelled aggressivity and reactivity/immaturity) that accounted for 56% of the total variance. Nine breed groups with different behavioural characteristics were generated by K-means cluster analysis. These groupings had similarities with the groupings presented in the USA and UK, e.g. of the seven breeds rated as high in aggressivity in this study, five were rated high in all three countries, the Miniature Schnauzer was rated high for aggressivity in Italy and the US, but the Yorkshire Terrier was rated high only in Italy. These results provide further evidence of the need for care when transposing breed behavioural advice or treatments between countries.
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    The behaviour test results of 1310 German shepherds and 797 Labrador retrievers, 450–600 days of age, were evaluated. The purpose was to investigate whether the behaviour tests, previously used at the Swedish Dog Training Centre, could be used to select dogs for different kinds of work and for breeding. Ten behavioural characteristics were scored based on the dogs' reactions in seven different test situations. All tests were conducted by one experienced person.Marked differences in mental characteristics were found between breeds and sexes, but particularly between various categories of service dogs. Regardless of differences in the behaviour profiles of these service categories, there were marked similarities between different categories of service dogs compared with dogs found to be unsuitable for training as service dogs. To interpret the data, an index value was created, based on the test results for each individual dog, and was found to be an excellent instrument for selecting dogs for different types of work.For both breeds the factor analysis resulted in four factors. In comparing the different characteristics, the same pattern was found in both breeds, with the exception of the characteristic prey drive, which seems to be irrelevant for Labrador retrievers. The conclusion is that a subjective evaluation of complex behaviour parameters can be used as a tool for selecting dogs suitable as service dogs. The results also show that the use and correct interpretation of behaviour tests can be enhanced by adjusting the results for each breed and planned service category.
  • Article
    Dogs show considerable variation in morphology, genetics and behaviour caused by long periods of artificial selection. This is evident in the large number of breeds we have today. Behavioural differences among breeds have often been regarded as remnants from past selection during the breeds’ origin. However, the selection in many breeds has, during the last decades, gone through great changes, which could have influenced breed-typical behaviour. In order to investigate this, breed differences were studied using data from a standardized behavioural test from 13,097 dogs of 31 breeds from the Swedish dog population. Based on the test results, breed scores were calculated for four behavioural traits: playfulness, curiosity/fearlessness, sociability and aggressiveness. These traits have previously been found to be stable and valid, and hence regarded as personality traits in the dog. The present results suggested large differences between breeds in all of the investigated traits, even though there were within-breed variations. No relationships between breed-characteristic behaviour and function in the breeds’ origins were found. Instead, there were correlations between breed scores and current use of the breeding stocks, which suggest that selection in the recent past has affected breed-typical behaviour. The breeds’ use in dog shows, the dominating use in general, was negatively correlated with all investigated traits, both in sires and in dams. In contrast, use in Working dog trials was positively correlated with playfulness and aggressiveness in sires. Thus, these results suggest that selection for dog show use is positively correlated with social and non-social fearfulness, and negatively with playfulness, curiosity in potentially threatening situations and aggressiveness, whereas selection for Working dog use is positively correlated with playfulness and aggressiveness. Furthermore, correlation analyses show that popular breeds have higher sociability and playfulness scores than less popular breeds, suggesting that a positive attitude towards strangers is an important characteristic of a functional pet dog and desirable by dog owners. This indicates that selection towards use in dog shows may be in conflict with pet dog selection. Furthermore, these results suggest that basic dimensions of dog behaviour can be changed when selection pressure changes, and that the domestication of the dog still is in progress. A standardized behavioural test, like the one used in this study, is suggested to be highly useful as a tool in dog breeding programs.
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    Heritability calculated for characteristics evaluated in behavioural tests can be used as a tool to select different kinds of service dogs. The evaluation was based on the test results of 1310 German shepherds and 797 Labrador retrievers. The heritability for all evaluated characteristics of the two breeds was significantly different from zero with the exception of the characteristics prey drive in Labrador retrievers.The test results for each characteristic were summarised to form an index value which simplified the interpretation of the test results. The heritability for this index value was 0.24 for both German shepherds and Labrador retrievers, a value that must be considered high as it included all tested parameters. The heritability was also calculated for the four factors derived from a factor analysis of the test results. Heritability estimates for these four factors were 0.15 to 0.32The results show that complex behavioural patterns in dogs can be subjectively evaluated by an experienced person and that no more than a few characteristics are needed in order to describe the differences between dogs.Breeding results in a German shepherd population at the Swedish Dog Training Centre (SDTC) improved a relatively short time after the initiation of basing the selection of breeding animals on the index value of each individual animal. German shepherds bred by the SDTC also had higher index values than privately bred dogs which shows the importance of a goal-oriented breeding programme with emphasis on service dog characteristics.Finally different ways in which to collect information about dog behaviour are discussed. It is suggested that a subjective evaluation of certain behaviour characteristics is preferred to a factual description of reactions.
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    Many companion dogs occupy a privileged position in our society, living closely with human caretakers who go to great lengths to provide for their needs and desires. Others fare less well, being abandoned or killed, many because they are believed to exhibit behaviour problems. The aim in this study was to investigate the frequency of potentially problematic behaviours experienced by a convenience sample of companion dog owners and to establish if the presence of these behaviours was associated with demographic variables, involvement in dog training activities and participation in other dog-human interactions. Potentially problematic behaviours were reported to occur by the 413 adult participants only infrequently, but fell into five factors; disobedience, unfriendliness/aggression, nervousness, anxiety/destructiveness and excitability. Each of these factors was associated with a number of owner and dog characteristics. Engagement in training activities was predictive of lower scores being obtained for many of the behaviours, as well as increased involvement in shared activities. Some of the behaviours, particularly the perceived friendliness of the dog, were also predictive of involvement in shared activities. This confirms that strategies designed to increase participation in dog training activities and promote canine sociability may have significant benefits for both companion dog owners and their dogs.