Content uploaded by Simona Cafazzo
Author content
All content in this area was uploaded by Simona Cafazzo on Jul 14, 2016
Content may be subject to copyright.
RESEARCH PAPER
Scent-Marking Behaviour in a Pack of Free-Ranging Domestic
Dogs
Simona Cafazzo*, Eugenia Natoli†& Paola Valsecchi*
* Dipartimento di Biologia Evolutiva e Funzionale, Universita
`degli Studi di Parma, Parma, Italy
†Azienda USL Roma D, Area Dipartimentale Sanita
`Pubblica Veterinaria, Rome, Italy
Correspondence
Simona Cafazzo, Dipartimento di Biologia
Evolutiva e Funzionale, Universita
`degli Studi
di Parma, via G.P. Usberti 11/A, 43100 Parma,
Italy, and via Gasperina 300, 00173 Rome,
Italy.
E-mail: simona.cafazzo@inwind.it
Received: June 20, 2011
Initial acceptance: September 16, 2011
Final acceptance: July 5, 2012
(M. Herberstein)
doi: 10.1111/j.1439-0310.2012.02088.x
Abstract
Most mammals scent-mark, and a variety of hypotheses have been put
forward to explain this behaviour. Most of our knowledge about scent
marking in domestic dogs comes from studies carried out on laboratory or
companion dogs, while few studies have been carried out on free-ranging
dogs. Here, we explored the functional significance of different scent-
marking behavioural patterns in a pack of free-ranging domestic dogs by
testing two non-exclusive hypotheses: the indirect territorial defence
and the dominance/threat hypotheses. Through direct observation, we
recorded the locations of dog scent marks (urination, defecation and
ground scratching) and information regarding the identity and posture of
the marking animal. We found evidence that markings are used by dogs
to form a ‘property line’ and to threaten rivals during agonistic conflicts.
Both males and females utilized scent marking to assert dominance and
probably to relocate food or maintain possession over it. Raised-leg urina-
tion and ground scratching probably play a role in olfactory and visual
communication in both males and females. Urinations released by
females, especially through flexed-leg posture, may also convey informa-
tion about their reproductive state. Finally, our observations suggest that
defecation does not play an essential role in olfactory communication
among free-ranging dogs and that standing and squat postures are associ-
ated with normal excretion. Our results suggest that many of the proposed
functions of marking behaviours are not mutually exclusive, and all
should be explored through detailed field and laboratory studies.
Introduction
Scent marking may be defined as the deposition of
odour by urination, defecation or the release of glan-
dular secretions (Kleiman 1966). Scent marking may
communicate detailed information about the scent
source, including social status, age, sex and reproduc-
tive state, group composition and individuality (e.g.
Ralls 1971; Gorman & Trowbridge 1989; Smith et al.
2001).
Most of our knowledge about scent marking in
domestic dogs (Canis lupus familiaris) comes from stud-
ies carried out on laboratory or companion dogs.
Urinary behaviour in domestic dogs is sexually dimor-
phic (Beach 1974; Ranson & Beach 1985). Adult
males urinate more frequently than do females and
are more likely than females to direct their urine at
specific targets (Martins & Valle 1948; Bekoff 1979a;
Ranson & Beach 1985). In addition, adult males typi-
cally raise a hind leg to urinate, while adult females
usually squat (Martins & Valle 1948; Beach 1974).
While urination clearly has the physiological function
to empty the bladder in both sexes, its role in scent
marking is thought to be more important in male dogs
than in female dogs (Martins & Valle 1948; Bekoff
1979a; Pal 2003). Indeed, Kleiman (1966) suggested
that female dogs mark with urine only around the
time they are in oestrus. Lisberg & Snowdon (2009,
2011) observed that, outside of oestrus, both high-
status males and females urinated, investigated urine
and countermarked more than same-sex dogs with
lower status. Moreover, males and females were
Ethology 118 (2012) 1–12 ©2012 Blackwell Verlag GmbH 1
Ethology
equally likely to countermark and investigate urine
(Lisberg & Snowdon 2011).
A study (Pal 2003) carried out using direct observa-
tion of urine-marking behaviour of free-ranging
domestic dogs further showed that urine marking was
displayed especially during territorial defence. Ques-
tions remain, however, regarding the function of
urine-marking behaviour and other scent-marking
behaviours, especially in relation to different social
classes within a pack.
In the current study, we explored the functional
significance of different scent-marking behaviours in
a pack of free-ranging domestic dogs living in a subur-
ban environment. We focused on three different
behavioural patterns of marking: urination, defeca-
tion and ground scratching. Scent marking by urina-
tion is a common practice in the family Canidae (e.g.
Scott & Fuller 1965; Kleiman 1966; Beach 1974; Fox
et al. 1975; Peters & Mech 1975; Dunbar 1977, 1978;
Bekoff 1979a; Wells & Bekoff 1981; Gese & Ruff 1997;
Allen et al. 1999; Pal 2003; Zub et al. 2003). Defeca-
tion and ground scratching can also serve as scent
marking in canids (e.g. wolves, Canis lupus: Peters &
Mech 1975; Barja & De Miguel 2004; coyote, Canis la-
trans: Wells & Bekoff 1981; Gese & Ruff 1997). Scats
often are left in conspicuous locations, such as on
rocks, plants and trail junctions, probably increasing
their effectiveness as visual and olfactory marks.
Ground scratching often occurs after defecation or
raised-leg urination, and it may serve as a chemical
signal and as a visual signal (Peters & Mech 1975;
Bekoff 1979b; Bekoff & Wells 1986). The chemical
component may involve either scent deposited from
interdigital glands or the dispersion of olfactory cues
from deposited urine or faeces (Peters & Mech 1975;
Fox & Cohen 1977). The visual component may
involve the act of ground scratching or the marks left
on the ground by such scratching (Kleiman & Eisen-
berg 1973; Bekoff 1979b). Ground scratching has not
been reported to follow squat urination and appears
to be performed predominantly by high-ranking
males (e.g. wolves: Peters & Mech 1975; coyotes: Gese
& Ruff 1997).
Our aim was to test two non-exclusive hypotheses
related to the function of scent marking by free-rang-
ing domestic dogs: the ‘indirect territorial defence
hypothesis’ and the ‘dominance/threat hypothesis’.
Animals may scent-mark for the purpose of defend-
ing food resources or territories, thereby reducing
fights with conspecifics (indirect territorial defence
hypothesis). Scent marks could work as a chemical
‘keep out sign’, which would deter or intimidate
intruders (Wyatt 2003), or scent marks allow an
intruder to identify the territory owner by matching
scent marks encountered in the territory with the
scent of individuals encountered (Gosling (1982).
Identification of the territory owner by the intruder
can help to avoid escalated conflicts between them
and thus is beneficial to both parties (Gosling &
Roberts 2001). If the costs of scent-marking the terri-
tory (in terms of time and energy and perhaps
increased vulnerability to predators) are lower than
the costs of attacking an intruder, scent marking
becomes an economic strategy for territorial defence
(Gosling 1982, 1986; Gosling & Roberts 2001).
According to both scent fence and scent-matching
mechanisms, scent marking should be concentrated
in areas where scent marks are most likely to be
encountered by an intruder, that is, along the territo-
rial boundary (Gosling 1982). Studies on different
mammalian taxa have supported this prediction (e.g.
wolves, Peters & Mech 1975; Rothman & Mech 1979;
Zub et al. 2003; tigers, Panthera tigris: Smith et al.
1989; marmots, Marmota spp.: Boero 1995; Blumstein
& Henderson 1996; blind mole-rats, Spalax ehrenbergi,
Zuri et al. 1997; coyote, Wells & Bekoff 1981; Gese &
Ruff 1997; Allen et al. 1999; ethiopian wolves, Canis
simensis, Sillero-Zubiri & Macdonald 1998).
Ralls (1971) hypothesized that social mammals tend
to mark frequently in any situation where they are
both intolerant of and dominant to other members of
the same species. In other words, they mark when
they are likely to attack another member of the same
species and are likely to win if they do attack. Such
behaviour may occur in territorial defence, but it will
not necessarily be restricted to territorial situations. In
fact, scent marking may function to communicate
dominance status among group members, thus reduc-
ing the need for costly fights. The dominance/threat
hypothesis has received support from several studies
on scent marking in mammals (e.g. Walther 1978;
Gosling & Roberts 2001; Miller et al. 2003; Bonanni
et al. 2007).
In this study, we aim to test the general prediction
that, if scent marking is used to communicate indirect
defence of territory (territorial defence hypothesis), dogs
would mark more frequently along the periphery of the
territory than in its interior (prediction 1a); moreover,
dogs are expected to mark more frequently at strategic
places, such as crossroads, where potential intruders are
most likely to encounter marks (prediction 1b).
According to the dominance/threat hypothesis, we
predicted that dominant animals would mark more
frequently than subordinate animals (prediction 2a).
We also may predict a positive correlation between
scent marking and aggressive behaviour (prediction
Ethology 118 (2012) 1–12 ©2012 Blackwell Verlag GmbH2
Scent-Marking Behaviour in Domestic Dogs S. Cafazzo, E. Natoli & P. Valsecchi
2b) and an increase in marking rate prior to or during
agonistic interactions than during non-agonistic
encounters (prediction 2c).
Finally, to confirm and extend the results of previ-
ous research, in this study, we also asked the follow-
ing questions: is the rate and posture of scent marking
influenced by sex of dogs? Do dogs scent-mark at
different rates in different social contexts and in differ-
ent places? Do female dogs use urination, ground
scratching and defecation to mark their territory or to
indicate dominance?
Methods
Study area
The research was carried out in a suburban environ-
ment situated in the south-western outskirts of Rome,
Italy (41°50′N, 12°23′W; elevation: about 60 m). The
study area was about 300 ha and was delimited to the
north, west and south by roads with heavy traffic and
to the east by cultivated areas.
The south-west sector was urbanized, although
not densely populated. The north-east sector was
mainly occupied by a nature reserve called ‘Tenuta
dei Massimi’.
The habitat in the reserve consists mainly of open
grasslands with interspersed wooded areas (see
Bonanni et al. 2010a for a more detailed description).
All dog packs had free access to virtually every part
of the study area. Nevertheless, the individuals of the
pack studied mainly frequented the area of the Natu-
ral Park where the dense vegetation of the wooded
areas offered good shelter for the animals, especially
for lactating females and their puppies. However, they
frequently approached a central road crossing the
study area, especially in the early morning, to feed
on the food (mainly meat from a slaughterhouse)
brought every day by volunteer dog caretakers.
Subjects
The dog pack studied belonged to a population of
about 100 adult animals inhabiting the study area. All
dogs were mediumsized to large mongrels, and there
was not a recognizable predominant breeding type
(Cafazzo 2007). With very few exceptions, dogs were
not socialized to humans although they were depen-
dent on humans for food provisioning. All dogs of the
pack were individually recognized by coat colour and
pattern, hair length and body size; they were sexed by
genital morphology (for a detailed description, see
Cafazzo et al. 2010 and Bonanni et al. 2010a,b).
Observations of this pack of dogs began in Mar.
2005. We had direct knowledge of age in dogs 2 yr
and younger. The ages of the remaining dogs were
assessed by estimating body size and general appear-
ance (e.g. white hair on the muzzle) as well as tooth
wear (e.g. Gier 1968) and eruption (Kirk 1977) during
capturing and immobilization procedure (see Cafazzo
et al. 2010 for details of age estimation). Dogs were
thus reliably classified as juveniles (6 mo to 1 yr),
subadults (1–2 yr) and adults (more than 2 yr). At
the start of the research, the dog pack consisted of 27
individuals. Owing to some deaths, births and disap-
pearances of some individuals during the period of
observation (see. Cafazzo et al. 2010 for a detailed
description), the pack size ranged from 25 to 42 indi-
viduals. The final statistical analysis was applied to 27
dogs: six adult males, five adult females, four subadult
males, one subadult female, six juvenile males and
five juvenile females.
Behavioural observations
The study began in Apr. 2005 and lasted until the end
of May 2006. Data were collected in three different
contexts: in the presence of food, when receptive
females were courted by males (and food was not
present), and in the absence of sources of competition
(i.e. when food was not present and outside the oes-
trus period) for a total of 282.53 h of recording. Data
collection was carried out following Altmann’s (1974)
methods: focal animal sampling method was used in
the absence of sources of competition, whereas the
focal subgroup sampling method was used in the
presence of food and receptive females.
For each urination, defecation and ground scratch-
ing performed by the focal animals, the type, time and
location of the behaviour were recorded. According to
Kleiman (1966), urinations were classified as scent-
marking events rather than eliminations; when urine
was released in small quantity following olfactory
investigation, it was orientated towards a conspicuous
substrate (plants, rocks, food, walls, car wheels, etc.),
elicited by familiar landmarks, novel odours or objects
and conspecifics marks (usually dogs marked just over
or some distance away from investigated spots). Uri-
nation posture recorded included: (1) raised-leg uri-
nation (one hind leg is raised off the ground, whether
or not there is a vertical target); (2) flexed-leg urina-
tion (the dog squats and elevates one hind leg slightly
forward or directly under the body); (3) standing uri-
nation (dogs do not squat but they stand with their
hind legs straight); (4) squat urination (the dog’s hind
part is lowered, the hind legs are bent under the body
Ethology 118 (2012) 1–12 ©©2012 Blackwell Verlag GmbH 3
S. Cafazzo, E. Natoli & P. Valsecchi Scent-Marking Behaviour in Domestic Dogs
and the urogenital region comes close to the ground).
We also recorded all events of defecation. Scats were
considered as true marking cues if they were depos-
ited following olfactory investigation at conspicuous
or raised sites (Kleiman 1966). Finally, we also
recorded all events of ground scratching, the alternate
movements of the legs through which the dog
scratches the ground, sending grass and earth flying
backward; it often occurs after raised-leg urination
and defecation (Rothman & Mech 1979; Zub et al.
2003).
The individual measure of behavioural patterns
was corrected for animal observation time because
the latter varied between individuals.
In the absence of sources of competition, we also
recorded the location of the focal animal using the
instantaneous- sampling method (60-s intervals)
(Altmann 1974). Thus, we could estimate for each
dog, the time spent in the following zones: along the
territorial boundaries, outside the territory, at feeding
sites, at denning sites and somewhere else in the
territorial area.
Home range and territory sizes
To analyse the spatial distribution of scent marking,
we calculated the sizes of the home range and of the
defended area by applying the minimum convex poly-
gon method (Southwood & Henderson 2000). Home
range was examined by plotting the locations of the
animals sighted covering fixed paths everyday twice
per day. We totalled 150.65 ±19.95 sightings for each
dog during 197 days of observation. Home range size
was estimated by drawing a convex polygon connect-
ing the outermost sightings and computing its area,
including inner buildings and fenced areas.
As a counterevidence of the territory size esteem,
the locations where pack members clearly won the
territorial fights against intruders were plotted. A ter-
ritorial fight was defined as an agonistic encounter
between pack members under study and dogs belong-
ing to neighbouring social packs. The pack members
were considered winning the agonistic encounter
when they managed to drive away the intruders.
Statistical analysis
By using data on the direction of submissive behav-
iour observed between dogs, we could arrange all
individuals belonging to the pack studied in a linear
dominance hierarchy whose details are described
elsewhere (Cafazzo et al. 2010).
To test the predictions generated from the indirect
territorial defence hypothesis, we expressed the indi-
vidual marking rate as the number of markings
recorded in each zone divided for the time spent in
the same zone. Then, we compared the individual
marking rate displayed along territorial boundaries (at
crossroads or not) with those displayed both outside
(about 100 m beyond the territorial boundaries) and
inside the territorial area. A scent mark was classified
as a boundary mark when it occurred within 50 m of
the territorial boundaries. We also compared the
hourly rate of scent markings observed at denning
sites, feeding sites (when no food was present) and
somewhere else to analyse the spatial distribution of
scent marking inside the territorial area. For these
analyses, we only considered the scent markings
recorded in the absence of sources of competition
(scent marking that occurred during territorial conflict
or in the presence of food or oestrus females were not
included) because the spatial distribution of scent
marking observed in the other two contexts (during
feeding and courting) was influenced by the location
of feeding sites and by movements of receptive
females.
As in this dog pack the dominance rank, the rate of
aggressive behaviour and the age of the individuals
were correlated (see Cafazzo et al. 2010), we applied
a principal components analysis (PCA) to replace the
mentioned three original variables with new uncorre-
lated component variables, linear combinations of the
original variables, called principal components or fac-
tors. Then, we tested the dominance/threat hypothe-
sis by correlating the hourly rate of marking
behaviours to the PCA components that explained
most of the variation in the data.
By comparing the rate of scent marking during an
agonistic conflict with the rate of marking during
non-agonistic interactions, we verified the relation
between marks and the aggressive motivation towards
other dogs. For this analysis, we separately considered
interactions between pack members and with dogs
from other packs.
We tested the hypotheses and investigated the
relationship between scent marking and sex and
social context using two-tailed non-parametric sta-
tistical tests [Wilcoxon signed-ranks test (t), Mann–
Whitney U-test, and Spearman rank correlation
(r
s
)], (STATISTICA 7.1 edition, StatSoft Italy s.r.l.
2005). Probability level for rejection of the null
hypothesis was set at p <0.05. Media and range
of behavioural patterns analysed are listed in
Table 1.
Ethology 118 (2012) 1–12 ©2012 Blackwell Verlag GmbH4
Scent-Marking Behaviour in Domestic Dogs S. Cafazzo, E. Natoli & P. Valsecchi
Results
Home range and territory
Although some subgroups sometimes covered longer
distances than other subgroups, pack members usu-
ally moved all together throughout the area, and indi-
viduals’ home ranges clearly overlapped within the
pack. The home range pack size was calculated as
xof
the individuals’ home ranges (N =27), and it aver-
aged 60.98 ±9.18 ha.
Territorial boundaries were assumed to be the
same for all members of the pack because it has
been observed that all pack members cooperated for
territorial and resources defence against intruders
from other neighbouring social packs (Bonanni et al.
2010a). Group territory size measured 22.14 ha and
its boundaries were inside the boundaries of the home
range. We defined the periphery of the territory as a
zone within 50 m of the territorial boundaries. This
periphery or boundary zone comprised about 21% of
the area of territory. All dens, resting and feeding sites
were located inside the territory and never in the
periphery.
Scent-marking behaviours: influence of sex and
context
We recorded a total of 782 urinations, 143 ground
scratches and 89 defecations during 282.53 h of
observation. Of 782, 589 were raised-leg, 52 flexed-
leg, 36 standing and 105 were squat urinations.
All squat and standing urinations were identified as
simple eliminations because they were never pre-
ceded by olfactory investigation, and they were
released on inconspicuous substrates (the ground).
For males, the incidence of marking was greater than
the incidence of simple elimination (Wilcoxon signed
rank test: t=1.00, z=3.46, n =16, p =0.0005). On
the contrary, females more frequently displayed
normal excretion than marking (t=7.00, z=2.09,
n=11, p =0.04).
Of 89 defecations recorded, only nine scats have
been considered to have a communicative function
because they were deposited following investigation
on raised sites (on rocks or plants); moreover, these
nine scats were deposited at crossroads. The incidence
of scent marking via urination was greater than
the incidence of scent marking via defecation.
The proportion of different postures displayed dur-
ing urination varied among the two sexes. Males
typically performed raised-leg urination (N =581;
94.01%), while they rarely displayed standing urina-
tion posture (N =36; 5.83%); only a juvenile male
was observed displaying a squat posture (N =1,
0.16%). Females rarely displayed raised-leg urination
(N =8, 4.88%), while they usually displayed flexed-
leg (N =52, 31.71%) and squat urination postures
(N =104; 63.42%).
The sex of dogs influences the frequency of scent
marking via urination: the marking rate via urination
was higher for males than for females (Mann–
Whitney U-test: U=42.5, n1 =16, n2 =11, p =
0.023). Females included a different representation of
age categories than males because only one subadult
female was observed. As this could skew the results,
we also compared only adult dogs when looking at
male vs. female marking, and we found again that
males marked via urination more frequently than
females (U=2.37, n1 =6, n2 =5, p =0.017).
The hourly rate of raised-leg urination displayed by
males was higher during feeding than in the presence
of receptive females; although the male raised-leg uri-
nation rate was higher in the absence of sources of
competition than in the presence of food, the differ-
ence was not significant (Table 2). Male raised-leg
urination rate was also higher in the absence of
sources of competition than in the presence of recep-
tive females (Table 2). Males ground-scratched more
frequently in the presence of food than in the pres-
ence of receptive females and even more frequently
in the absence of sources of competition (Table 2).
Standing urination was never observed in the pres-
ence of receptive females and its hourly rate did not
differ across the other two social contexts (Table 2).
Females displayed flexed-leg urination more fre-
quently during their oestrus than in the presence of
food and in the absence of sources of competition;
moreover, the hourly rate of flexed-leg urination was
higher in the presence of food than in the absence of
Table 1: Media and range of hourly rates of behavioural patterns
considered
N Media Minimum Maximum
Urination marking displayed
by males
16 0.72 0.13 2.67
Urination marking displayed
by females
11 0.27 0.00 0.72
Ground-scratching displayed
by all dogs
27 0.11 0.00 0.48
Standing urination displayed
by males
16 0.07 0.00 0.22
Squat urination displayed
by females
11 0.47 0.12 0.71
Defecation displayed by
all dogs
27 0.11 0.03 0.19
Ethology 118 (2012) 1–12 ©©2012 Blackwell Verlag GmbH 5
S. Cafazzo, E. Natoli & P. Valsecchi Scent-Marking Behaviour in Domestic Dogs
sources of competition (Table 3). Squat urination
hourly rate did not differ among the three social con-
texts (Table 3). All ground scratchings and raised-leg
urinations performed by females were recorded in the
absence of sources of competition.
Test of predictions
Prediction 1
Males scent-marked more frequently along territorial
boundaries and in the interior of the territory than
in rest of their home range (Table 4), while females
were never observed marking outside the territorial
area.
The hourly rate of scent marking via urination
along territorial boundaries vs. the interior was
influenced by the sex of dogs. Males scent-marked
via raised-leg urination along territorial boundaries
at a higher frequency compared with the interior
(Table 4) and mainly at the crossroads along territo-
rial borders (t=5.00, z=2.98, n =16, p =0.003). In
contrast, females scent-marked via flexed-leg urina-
tion more frequently inside the territorial area than
along the boundaries (Table 4). All events of raised-
leg urination by females (N =8) were observed along
the periphery at the crossroads. Also males scent-
marked via raised-leg urination mainly at the cross-
roads along territorial borders (t=5.00, z=2.98,
n=16, p =0.003). Inside the territory, females scent-
marked more frequently at denning sites than at feed-
ing sites and somewhere else (Table 5), even if the
urine-marking rate was higher at feeding sites than
somewhere else the difference was not significant
(Table 5). On the contrary, the urine-marking rate of
males was higher at feeding sites than denning sites
and somewhere else (Table 5), while it did not differ
between denning sites and somewhere else (Table 5).
Males ground-scratched along territorial boundaries
at a higher frequency compared with the inside
(t=0.00, z=2.67, n =16, p =0.007) and outside of
territory (t=0.00, z=2.64, n =16, p =0.006), but no
difference was found between the ground-scratching
rate recorded inside and outside the territory (t=
11.00, z=1.21, n =16, p =0.36). Females scratched
the ground only along territorial boundaries following
raised-leg urination.
Scent marking via defecation was displayed only by
the alpha male along territorial boundaries. Of 11
events of defecation displayed by the alpha male, 9
(81.82%) were deposited along territorial boundaries
at crossroads and mainly on conspicuous or high
substrates (rocks and plants; N =7, 77.78%).
All elimination events and the 80 events of defeca-
tions identified as normal excretions were randomly
distributed: along territorial boundaries, in the inte-
rior of the territory and in the rest of the home range
(Table 6).
Prediction 2
For males, the first factor of the PCA alone explained
86.66% of the total variance in the data (Fig. 1a):
Table 2: We used the Wilcoxon signed-ranks test to compare the hourly rates of raised-leg urination, ground scratching and standing urination
performed by males, among three competitive contexts: in the absence of sources of competition, in the presence of food and during courting of
receptive females
Raised-leg urination Ground scratching Standing urination
tzptzptz p
Absence of sources of competition vs. presence of food (n =16) 49 0.98 0.32 5.00 2.07 0.04 36 0.66 0.50
Presence of food vs. during courting (n =16) 0.00 3.40 0.001 1.00 2.38 0.02 a a a
Absence of sources of competition vs. during courting (n =16) 0.00 3.40 0.001 0.00 2.67 0.01 a a a
a
Standing urination was never observed in the presence of receptive females.
Table 3: We used the Wilcoxon signed-ranks test to compare the hourly rates flexed-leg urination and squat urination performed by females among
three competitive contexts: in the absence of any sources of competition, in the presence of food and during courting by males
Flexed-leg urination Squat urination
tzptzp
Absence of sources of competition vs. presence of food (n =11) 3.00 2.10 0.04 15.00 1.60 0.11
Presence of food vs. during courting (n =7) 0.00 2.37 0.002 11.00 0.50 0.51
Absence of sources of competition vs. during courting (n =7) 0.00 2.36 0.002 5.00 1.52 0.13
Ethology 118 (2012) 1–12 ©2012 Blackwell Verlag GmbH6
Scent-Marking Behaviour in Domestic Dogs S. Cafazzo, E. Natoli & P. Valsecchi
males characterized by high negative scores on this
factor were high-ranking older individuals who
displayed a higher frequency of aggressive behaviour.
The second factor of the PCA explained just 12.01%
of the variance (Fig. 1a).
The first factor was negatively correlated with both
urine scent-marking hourly rate (r
s
=0.95, n =16,
p=0.000001) and ground-scratching hourly rate
(r
s
=0.67, n =16, p =0.005) displayed by males.
Therefore, high-ranking old dogs were more aggres-
sive and displayed scent-marking behavioural pat-
terns more frequently than juveniles or subadults.
For females, the first factor of the PCA alone
explained 73.90% of the total variance in the data
(Fig. 1b); the second factor of the PCA explained
24.48% of the variance. Thus, females characterized
by high negative scores for the first factor were high-
ranking old individuals who displayed a higher fre-
quency of aggressive behaviour. Females with high
negative scores on the second factor displayed a
higher frequency of aggressive behaviour (Fig. 1b).
Moreover, these dogs were also younger, on average,
than those that had high negative scores on the first
component. The rate of scent marking via urination
displayed by females was negatively correlated to the
first factor (r
s
=0.88, n =11, p =0.0004), but it
was not correlated to the second factor (r
s
=0.28,
n=11, p =0.41).
Males scent-marked via urination more frequently
during agonistic than non-agonistic interactions with
both other pack members and dogs from other packs
(Table 7). Moreover, the rate of marking during ago-
nistic interactions with pack members was lower than
during agonistic interactions with dogs from other
packs (t=0.00, z=3.30, n =16, p =0.001). Females
also scent-marked via urinations (both flexed-leg and
raise-leg urinations) more often during agonistic than
non-agonistic interactions with both other pack mem-
bers and dogs from other packs (Table 7). However,
for them, the rate of marking during agonistic interac-
tions with pack members and with dogs from other
packs did not differ (t=5.00, z=0.67, n =11,
p=0.5). Finally, the rate of ground scratching was
higher during agonistic than non-agonistic interac-
tions with both pack members and dogs from other
packs (Table 7). Dogs ground-scratched at higher rate
Table 4: We used the Wilcoxon signed-ranks test to compare the hourly rate of raised-leg urination displayed by males (n =16) and the hourly rate
of flexed-leg urination displayed by females (n =11) outside and inside of the territory and along territorial boundaries
Raised-leg urination (n =16) Flexed-leg urination (n =11)
tzptzp
Outside of the territory vs. territorial boundaries 0.00 3.52 0.0004
Territorial boundaries vs. inside of the territory 0.00 3.51 0.0004 0.00 2.20 0.03
Outside of the territory vs. inside of the territory 1.00 2.85 0.0004
Table 5: We used the Wilcoxon signed-ranks test to compare the hourly rate of urine marking displayed by males (n =16) and females (n =11)
inside the territory at different sites: denning sites, feeding sites and somewhere else
Raised-leg urination (n =16) Flexed-leg urination (n =11)
tzptzp
Denning sites vs. feeding sites 0.00 2.98 0.003 0.00 2.22 0.02
Denning sites vs. somewhere else 10.00 1.12 0.26 0.00 2.18 0.04
Feeding sites vs. somewhere else 0.00 2.91 0.004 0.00 1.83 0.07
Table 6: We used the Wilcoxon signed-ranks test to compare the hourly rate of standing urination displayed by males (n =16), squat urination
displayed by females (n =11) and defecation displayed by all dogs (n =27) outside and inside of the territory and along territorial boundaries
Standing urination (n =16) Squat urination (n =11) Defecation (n =27)
tzptzptzp
Outside of the territory vs. territorial boundaries 13.00 0.17 0.87 7.00 1.18 0.24 103.00 0.76 0.45
Territorial boundaries vs. inside of the territory 3.00 1.86 0.07 30.00 0.27 0.79 135.00 1.30 0.19
Outside of the territory vs. inside of the territory 10.00 1.12 0.26 11.00 1.95 0.06 158.00 0.45 0.66
Ethology 118 (2012) 1–12 ©©2012 Blackwell Verlag GmbH 7
S. Cafazzo, E. Natoli & P. Valsecchi Scent-Marking Behaviour in Domestic Dogs
during agonistic interaction with dogs from other
pack than with pack members (t=0.00, z=3.06,
n=27, p =0.002).
Discussion
Indirect territorial defence hypothesis
Our results provide evidence that scent marking by
free-ranging domestic dogs may serve as demarcation
of territorial boundaries according to the territorial
defence hypothesis. Male dogs, as other species of
canids (wolves: Harrington 1981; Asa et al. 1985a,b,
1990; Harrington & Asa 2003; Peters & Mech 1975;
Zub et al. 2003; coyotes: Wells & Bekoff 1981; Gese &
Ruff 1997; Allen et al. 1999), appear to delimit their
territory by means of urine and scratching. Dogs, as
wolves (Zub et al. 2003) and coyotes (Wells & Bekoff
1981), were observed scent-marking especially at
crossroads, where the intrusion of other dogs from
neighbouring packs might be more likely. Neverthe-
less, whether scents alone (i.e. the scent-fence
hypothesis) or the matching between scents and sig-
nallers (territory owners; i.e. the scent-matching
hypothesis) would keep out intruders needs to be
explored. Moreover, as we did not study the scent-
marking distribution pattern of neighbouring dog
packs, we cannot exclude that the concentration of
scent marking along territorial boundaries could
also be caused by dogs countermarking unfamiliar
conspecifics’ urine; this does not necessarily indicate
territory defence as the main function of urine
marking.
Wolves have been observed depositing and accu-
mulating scats at strategic sites (e.g. crossroads, con-
spicuous substrates), probably to increase their
effectiveness as visual and olfactory marks (Barja
et al. 2004). In our dog pack, only the alpha male was
observed scent-marking by means of faeces at cross-
roads, but a real accumulation of scats in this place
has never been observed. As group members collec-
tively defended the territory in agonistic encounters
with intruders, if scats functioned in territorial
defence, a group-wide (rather than ‘alpha only’) use
of scats along boundaries, especially at crossroads,
should be expected. It is not clear why it was per-
formed only by the alpha male.
Contrary to males, female dogs scent-marked
mainly in the interior of the territory. Nevertheless,
the findings that both females and males scent-
marked inside the territory suggest that information
was also conveyed via these internal marks.
Dominance/threat hypothesis
Our results are also consistent with the dominance/
threat hypothesis (Ralls 1971). High-ranking individ-
uals are more likely to display marking behaviour.
(a)
(b)
Fig. 1: Principal components analysis (PCA) results for males (a) and
females (b). The circles shows the correlation between the original vari-
ables and the two new components. (a) Pearson correlations between
factor 1 of the PCA and the original variables were: 0.95 (Age), 0.97
(Hierarchical rank), 0.86 (Rate of aggressive behaviour). Pearson corre-
lations between factor 2 and the original variables were: 0.27 (Age),
0.17 (Hierarchical rank), 0.50 (Rate of aggressive behaviour). (b) Pear-
son correlations between factor 1 of the PCA and the original variables
were: 0.90 (Age), 0.98 (Hierarchical rank), 0.68 (Rate of aggressive
behaviour). Pearson correlations between factor 2 and the original vari-
ables were: 0.42 (Age), 0.12 (Hierarchical rank), 0.74 (Rate of aggres-
sive behaviour).
Ethology 118 (2012) 1–12 ©2012 Blackwell Verlag GmbH8
Scent-Marking Behaviour in Domestic Dogs S. Cafazzo, E. Natoli & P. Valsecchi
Moreover, the frequency of scent marking was clearly
related to aggressive behaviour given that dogs
marked more frequently during agonistic interactions
than non-agonistic interactions between both pack
mates and dogs from other packs. A higher frequency
of territorial marking behaviour performed by high-
ranking individuals was also found in wolves (Mech
1999, 2000; Peterson et al. 2002) and coyotes (Gese &
Ruff 1997), as well as in high-status companion dogs
(Lisberg & Snowdon 2011).
Males urinated raising a leg at higher rates during
conflicts with other dogs, especially against intruders
and along territorial boundaries; females displayed
raised-leg urinations only along territorial boundaries
and more often during agonistic interactions with
intruders. These findings support the idea that this
behavioural pattern may serve both as a mechanism
of border enforcement (as suggested for other species
of canids: coyotes, Gese & Ruff 1997; wolves, Barja
et al. 2004) and as a visual cue to display dominance
or threat to opponents.
The functions of ground scratching
Males scratched the ground mainly after raised-leg
urinations along territorial boundaries; females were
observed scratching the ground only after raised-leg
urinations. Both of them displayed this behaviour,
especially during territorial conflicts. Finally, high-
ranking older individuals scratched the ground more
often than low-ranking younger dogs. These results
highlight the role of ground scratching as a display of
social dominance and as a means of territorial
defence. In fact, scents dispersed during ground
scratching may identify the social status of the signal-
ler and inform intruders of territorial boundaries.
Therefore, from our results, ground scratching in
domestic dogs appears to be an additional visual mark
to reinforce the olfactory cue of urine marking and to
intimidate opponents.
Influence of sex
The rate of marking was higher in males than in
females as observed in other studies carried out on
free-ranging dogs (Bekoff 1979a; Pal 2003), laboratory
(Dunbar 1977, 1978) and companion dogs (Lisberg &
Snowdon 2011) as well as on wolves (Mech 2006).
Moreover, older individuals marked at higher rates
than younger as reported in previous studies on
domestic dogs (Wirant & McGuire 2004; Wirant et al.
2007) as well as on wolves (Sillero-Zubiri & Macdon-
ald 1998) and coyotes (Gese & Ruff 1997). Actually,
adults dominated over subadult dogs and subadults
dominated over juveniles; males dominated over
females in each age classes (see Cafazzo et al. 2010).
Therefore, high-ranking adult males scent-marked
more frequently than other age/sex categories.
Raised-leg urination posture was exhibited by males
and rarely by females, which used mainly flexed-leg
urination posture. Our results suggest that different
postures might be correlated with certain behaviours
and contexts: flexed-leg urination appeared to be
associated with reproductive state and with the
defence of dens and food; raised-leg urination seems
to be mainly associated with aggression and demarca-
tion of territorial boundaries in both males and
females.
Influence of the context
In free-ranging dogs in India, a higher rate of scent
marking has been reported during the oestrus period
(Pal 2003). Although we observed females scent-
marking more often during oestrus, males scent-
marked mainly during feeding and in the absence of
sources of competition than in the presence of
receptive females. Agonistic interactions among
male dogs were more frequent in the presence of
receptive females than in the absence of sources of
competition and even more frequent in the presence
Table 7: We used the Wilcoxon signed-ranks test to compare the hourly rate of urine marking displayed by males (n =16) and by females (n =11)
and the hourly rate of ground scratching by all dogs (n =27) during agonistic and non-agonistic interactions with both other pack members and dogs
from other packs
Urine marking by males
(n =16)
Urine marking by
females (n =11)
Ground scratching
(n =27)
tzptzptzp
Agonistic interactions vs. non-agonistic interactions with other
pack members
0.00 3.52 0.0004 2.50 2.17 0.03 0.00 2.93 0.003
Agonistic interactions vs. non-agonistic interactions with dogs
from other packs
2.5 3.14 0.001 0.00 2.67 0.008 0.00 3.29 0.001
Ethology 118 (2012) 1–12 ©©2012 Blackwell Verlag GmbH 9
S. Cafazzo, E. Natoli & P. Valsecchi Scent-Marking Behaviour in Domestic Dogs
of food (see Cafazzo et al. 2010). Although scent
marking is linked to aggressive interactions and the
rate of aggressions was high in the presence of
receptive females, males marked more frequently in
the absence of sources of competition and in the
presence of food than in the presence of receptive
females. In the absence of sources of competition,
dogs marked mainly along territorial boundaries;
therefore, the use of scent marking during inter-
group competition for territorial defence might have
a predominant role with respect to the use of scent
marking during competition for receptive females.
As for males, we observed a higher level of competi-
tion among females in the presence of food than in
the absence of sources of competition (see Cafazzo
et al. 2010). The higher frequency of scent marking
by both males and females during feeding may be
linked to the high level of competition for food
among pack members. Moreover, dogs often marked
on food after taking their meals and before leaving
the feeding site, also when sufficient food still
remained there. According to Pal (2003), it may be
hypothesized that scent marking may facilitate relo-
cation of food resources, as also observed in red
foxes, wolves and coyotes (Henry 1977; Harrington
1981, 1982); although from our data, we cannot
exclude that dog scent marking during feeding in
order to guard this resources instead.
Scent marking by females during their oestrus likely
functions, at least in large part, as a declaration of
their reproductive state (Beach & Gilmore 1949;
Eisenberg & Kleiman 1972; Fox & Cohen 1977; Pal
2003); on the other hand, the lowest frequency of
marking behaviour displayed by males in the presence
of receptive females might suggest their scent marks
are not directly associated with the courtship of
females, in contrast to what reported by Pal (2003).
Nevertheless, we cannot exclude that females gain
information about the quality of potential mates from
the quality of their odours released by markings, as
observed in other species (Brown & Macdonald 1985;
Rich & Hurst 1998, 1999).
Conclusion
These results provide further support for the idea that
free-ranging dogs still retain some features of the
social organization and communication of their
wild ancestor, the wolf, that evolved before domesti-
cation (Bonanni et al. 2010a, b; Cafazzo et al. 2010;
Bonanni et al. 2011). They also remind us that many
of the proposed functions of marking behaviours are
not mutually exclusive, and all should be explored
through detailed field and laboratory studies of all
types of marks.
Acknowledgements
We are grateful to the caretakers, Annamaria Andre-
ozzi and Mirella De Paolis, for their availability to
answer all our questions about dogs and to Dr Gius-
eppe Cariola working at the Veterinary Hospital of
Rome for his help and assistance during the study.
Special thanks are due to Mark Bekoff for providing
relevant literature and to Roberto Bonanni who criti-
cally read the drafts of this paper and improved its
quality. Finally, we are thankful to the editor and the
referees whose suggestions improved the manuscript.
This research was partially funded by the Department
of Education, University and Research of Italian
Government (PRIN 2004 to P.V.), and by University
of Parma (FIL 2005 to P.V.).
Literature cited
Allen, J. J., Bekoff, M. & Crabtree, R. L. 1999: An observa-
tional study of coyote (Canis latrans) scent-marking and
territorialiy in Yellowstone National Park. Ethology 105,
289—302.
Altmann, J. 1974: Observational study of behavior:
sampling methods. Behaviour 48, 227—265.
Asa, C. S., Peterson, E. K., Seal, U. S. & Mech, L. D. 1985a:
Deposition of anal-sac secretions by captive wolves
(Canis lupus). J. Mammal. 66,89—93.
Asa, C. S., Mech, L. D. & Seal, U. S. 1985b: The use of
urine, faeces and anal-secretions in scent-marking by a
captive wolf (Canis lupus) pack. Anim. Behav. 33,
1034—1036.
Asa, C. S., Mech, L. D., Seal, U. S. & Plotka, E.D. 1990:
The influence of social and endocrine factors on urine-
marking by captive wolves (Canis lupus). Horm. Behav.
24, 497—509.
Barja, I. & De Miguel, F. J. 2004: Variation in stimulus,
seasonal context and response to urine marks by
captive Iberian wolves (Canis lupus signatus). Acta Ethol.
7,51—57.
Barja, I., De Miguel, F. J. & Ba
´rcena, F. 2004: Importance
of the crossroads in faecal marking behaviour of the
wolves (Canis lupus). Naturwissenschaften 91, 489—
492.
Beach, F. A. 1974: Effects of gonadal hormones on urinary
behavior in dogs. Physiol. Behav. 12, 1005—1013.
Beach, F. A. & Gilmore, R. W. 1949: Responses of male
dogs to urine from females in heat. J. Mammal. 30,
391—392.
Bekoff, M. 1979a: Scent-marking by free-ranging domestic
dogs. Biol. Behav. 4, 123—139.
Ethology 118 (2012) 1–12 ©2012 Blackwell Verlag GmbH10
Scent-Marking Behaviour in Domestic Dogs S. Cafazzo, E. Natoli & P. Valsecchi
Bekoff, M. 1979b: Ground-scratching by male domestic
dogs: a composite signal. J. Mammal. 60, 847—848.
Bekoff, M. & Wells, M. C. 1986: Social ecology and
behavior of coyotes. Adv. Stud. Behav. 16, 251—338.
Blumstein, D. T. & Henderson, S. J. 1996: Cheek-rubbing
in golden marmots (Marmota caudata aurea). J. Zool.
238, 113—123.
Boero, D.L. 1995: Scent-deposition behaviour in alpine
marmots (Marmota marmota L.): its role in territorial
defence and social communication. Ethology 100,
26—38.
Bonanni, R., Cafazzo, S., Fantini, C. & Natoli, E. 2007:
Relationship between dominance rank and territorial
marking behaviour in an urban feral domestic cat
colony. Hystrix It. J. Mamm. 2, 298.
Bonanni, R., Valsecchi, P. & Natoli, E. 2010a: Pattern of
individual participation and cheating in conflicts
between groups of free-ranging dogs. Anim. Behav. 79,
957—968.
Bonanni, R., Cafazzo, S., Valsecchi, P. & Natoli, E. 2010b:
Effect of affiliative and agonistic relationships on leader-
ship behaviour in free-ranging dogs. Anim. Behav. 79,
981—991.
Bonanni, R., Natoli, E., Cafazzo, S. & Valsecchi, P. 2011:
Free-ranging dogs assess the quantity of opponents in
intergroup conflicts. Anim. Cog. 14, 103—115.
Brown, R. E. & Macdonald, D. W. 1985: Social Odours in
Mammals. Vol. 1 & 2. Clarendon Press, Oxford.
Cafazzo, S. 2007: Dinamiche sociali in un gruppo di
cani domestici (Canis lupus familiaris) liberi in
ambiente suburbano. Ph.D. thesis, Univ. of Parma,
Parma, Italy.
Cafazzo, S., Valsecchi, P., Bonanni, R. & Natoli, E. 2010:
Dominance in relation to age, sex and competitive con-
texts in a group of free-ranging domestic dogs. Behav.
Ecol. 21, 443—455.
Dunbar, I. 1977: Olfactory preferences in dogs: the
response of male and female beagles to conspecific
odours. Behav. Biol. 20, 471—481.
Dunbar, I. 1978: Olfactory preferences in dogs: the
response of male and female beagles to conspecific
urine. Behav. Biol. 3, 273—286.
Eisenberg, J. F. & Kleiman, D. G. 1972: Olfactory
communication in mammals. Annu. Rev. Ecol. Syst.
3,1—31.
Fox, M. W. & Cohen, J. A. 1977: Canid communication.
In: How Animals Communicate (Sebeok, T., ed.). Indi-
ana Univ. Press, Bloomington, Indiana, pp. 728—748.
Fox, M. W., Beck, A. M. & Blackman, E. 1975: Behaviour
and ecology of a small group of urban dogs (Canis famili-
aris). Appl. Anim. Ethol. 1, 119—137.
Gese, E. M. & Ruff, R. L. 1997: Scent marking by coyotes,
Canis latrans: the influence of social and ecological
factors. Anim. Behav. 54, 1155—1166.
Gier, H. 1968: Coyotes in Kansas. Agric. Expt. Sta. Bull.
Kansas State Univ. 393,1—118.
Gorman, M. L. & Trowbridge, B. J. 1989: The role of odor
in the social lives of carnivores. In: Carnivore Behav-
iour, Ecology and Evolution (Gittleman, J. L., ed.). Cor-
nell Univ. Press, Cambridge, UK, pp. 57—88.
Gosling, L. M. 1982: A reassessment of the function of
scent marking in territories. Zeitschrift fu
¨r Tierpsycholo-
gie 60,89—118.
Gosling, L. M. 1986. Economic consequences of scent
marking in mammalian territoriality. In: Chemical Sig-
nals in Vertebrates (Duvall, D. & Muller-Schwarze, D.,
eds.). Plenum Press, New York, NY, pp. 385—395.
Gosling, L. M. & Roberts, S. C. 2001: Scent-marking by
male mammals: cheat-proof signals to competitors and
mates. Adv. Study Behav. 30, 169—217.
Harrington, F. H. 1981: Urine-marking and caching
behavior in the wolf. Behaviour 76, 280—288.
Harrington, F. H. 1982: Urine marking at food caches in
captives coyotes. Can. J. Zool. 60, 776—782.
Harrington, F. H. & Asa, C. S. 2003: Wolf communication.
In: Wolves: Behavior, Ecology and Conservation (Mech,
L. D. & Boitani, L., eds). Univ. of Chicago Press,
Chicago, IL, pp. 66—103.
Henry, J. D. 1977: The use of urine marking in the
scavenging behavior of the red fox (Vulpes vulpes).
Behaviour 61,82—105.
Kirk, R. W. (ed.) 1977: Current veterinary therapy. Vol.
VI: Small Animal Practice. W.B. Saunders Co, Philadel-
phia, PA.
Kleiman, D. 1966: Scent marking in the canidae. Symp.
Zool. Soc. Lond. 18, 167—177.
Kleiman, D. & Eisenberg, J. F. 1973: Comparisons of canid
and felid social systems from an evolutionary perspec-
tive. Anim. Behav. 21, 637—659.
Lisberg, A. E. Snowdon, C. T. 2009: The effects of sex,
gonadectomy and status on investigation patterns of
unfamiliar conspecific urine in domestic dogs, Canis
familiaris. Anim. Behav. 77, 1147—1154.
Lisberg, A. E. Snowdon, C. T. 2011: Effects of sex, social
status and gonadectomy on countermarking by domes-
tic dogs, Canis familiaris. Anim. Behav. 81, 757—764.
Martins, T. & Valle, J. R. 1948: Hormonal regulation of the
micturition behavior of the dog. J. Comp. Physiol.
Psychol. 41, 301—311.
Mech, L. D. 1999: Alpha status, dominance, and division
of labor in wolf packs. Can. J. Zool. 77, 1196—1203.
Mech, L. D. 2000: Leadership in wolf, Canis lupus, packs.
Can. Field- Nat. 114, 259—263.
Mech, L. D. 2006: Urine-marking and ground-scratching
by free-ranging Arctic wolves, Canis lupus arctos,in
Summer. Can. Field- Nat. 120, 466—470.
Miller, K. E., Laszlo, K. & Dietz, J. M. 2003: The role of
scent marking in the social communication of wild
Ethology 118 (2012) 1–12 ©©2012 Blackwell Verlag GmbH 11
S. Cafazzo, E. Natoli & P. Valsecchi Scent-Marking Behaviour in Domestic Dogs
golden lion tamarins, Leontopithecus rosalia. Anim.
Behav. 65, 795—803.
Pal, S. K. 2003: Urine marking by free-ranging dogs (Canis
familiaris) in relation to sex, season, place and posture.
Appl. Anim. Behav. Sci., 80,45—59.
Peters, R. P. & Mech, L. D. 1975: Scent-marking in wolves.
Am. Sci. 63, 628—637.
Peterson, O. R., Jacobs, A. K., Drummer, T. D., Mech, L. D.
& Smith, D. W. 2002: Leadership behaviour in relation
to dominance and reproductive status in gray wolves,
Canis lupus. Can. J. Zool. 80, 1405—1412.
Ralls, K. 1971: Mammalian scent marking. Science 171,
443—449.
Ranson, E. & Beach, F. A. 1985: Effects of testosterone on
ontogeny of urinary behavior in male and female dogs.
Horm. Behav. 19,36—51.
Rich, T. J. & Hurst, J. L. 1998: Scent marks as reliable sig-
nals of the competitive ability of mates. Anim. Behav.
56, 727—735.
Rich, T. J. & Hurst, J. L. 1999: The competing countermarks
hypothesis: reliable assessment of competitive ability by
potential mates. Anim. Behav. 58,1027—1037.
Rothman, R. J. & Mech, L. D. 1979: Scent-marking in
lone wolves and newly formed pairs. Anim. Behav. 27,
750—760.
Scott, J. P. & Fuller, J. L. 1965: Genetics and the Social
Behaviour of the Dog. Univ. of Chicago Press, Chicago, IL.
Sillero-Zubiri, C. & Macdonald, D. W. 1998: Scent-mark-
ing and territorial behaviour of Ethiopian wolves Canis
simensis. J. Zool. 245, 351—361.
Smith, J. L. D., McDougal, C. & Miquelle, D. 1989: Scent
marking in free-ranging tigers, Panthera tigris. Anim.
Behav. 37,1—10.
Smith, T. E., Tomlinson, A. J., Mlotkiewicz, J. A. &
Abbott, D. H. 2001: Female marmoset monkeys (Cal-
lithrix jacchus) can be identified from the chemical
composition of their scent marks. Chem. Senses 26,
449—458.
Southwood, T. R. E. & Henderson, P. A. 2000: Ecological
Methods. Blackwell Science, Oxford.
Walther, F. R. 1978: Mapping the structure and the mark-
ing system of a territory of the Thomson’s gazelle. E.
Afr. Wildl. J. 16, 167—176.
Wells, M. C. & Bekoff, M. 1981: An observational study of
scent-marking in coyotes, Canis latrans. Anim. Behav.
29, 332—350.
Wirant, S. C. & McGuire, B. 2004: Urinary behaviour of
female domestic dogs (Canis familiaris): influence of
reproductive status, location, and age. Appl. Anim.
Behav. Sci. 85, 335—348.
Wirant, S. C., Halvorsen, T. & McGuire, B. 2007: Prelimin-
ary observations on the urinary behaviour of female
Jack Russell Terriers in relation to stage of the oestrous
cycle, location, and age. Appl. Anim. Behav. Sci. 106,
161—166.
Wyatt, T. D. 2003: Pheromones and Animal Behaviour.
Cambridge Univ. Press, Cambridge, UK,
pp. 87—101.
Zub, K., Theuerkauf, J., Jedrzejewski, W., Jedrzejewska,
B., Schmidt, K. & Kowalczyk, R. 2003: Wolf pack terri-
tory marking in the Bialowieza primeval forest
(Poland). Behaviour 140, 635—648.
Zuri, I., Gazit, I. & Terkel, J. 1997: Effect of scent-
marking in delaying territorial invasion in the
blind mole-rat Spalax ehrenbergi. Behaviour 134,
867—880.
Ethology 118 (2012) 1–12 ©2012 Blackwell Verlag GmbH12
Scent-Marking Behaviour in Domestic Dogs S. Cafazzo, E. Natoli & P. Valsecchi
