Isabel BARJA*, Francisco Javier de MIGUEL
Departamento Biología, Unidad Zoología, Universidad Autónoma de Madrid, 28049 Madrid, Spain
*e-mail: firstname.lastname@example.org (corresponding author)
CHEMICAL COMMUNICATION IN LARGE CARNIVORES:
URINE-MARKING FREQUENCIES IN CAPTIVE TIGERS
POLISH JOURNAL OF ECOLOGY
(Pol. J. Ecol.)
ABSTRACT: Environmental and social pres-
sures can result in interspecies differences in mark-
ing behaviours. There is a strong relationship be-
tween marking behaviour and the environment.
Therefore, closely related species that show behav-
ioural differences in the wild may have different
scent marking strategies. We conducted a compara-
tive study of the urine-marking behaviours of tigers
and lions in captivity (Madrid Zoo, open enclosures
of 514 m2 and 730 m2 respectively, observations of 8
animals for each species). These two closely related
species have different natural habitats. We observed
interspecific differences in the rates, seasonal varia-
tions, and durations of the urine-marking acts.
The marking rate was higher in tigers, which also
showed seasonal variations not observed in lions.
The duration of urine marking was lower in tigers
than in lions. These differences seem to correspond
to differences between tigers and lions in terms of
their natural habitats (forest areas vs open areas),
social organizations (solitary vs social), and repro-
ductive biology patterns (seasonal polyoestrous vs
KEY WORDS: captivity, chemical communi-
cation, lions, tigers, urine marking
In order to increase their fitness, wild
animals must defend valuable resources, such
as food, mates, and territories, against their
competitors. The defence of resources can
involve aggression, threatening, or marking
behaviours, which are used to label the envi-
ronment with visual or scent marks. Mark-
ing strategies depend upon the animal’s life-
style, social requirements, and the physical
features of their environment. In mammals,
particularly nocturnal or crepuscular mam-
mals, chemicals are often used to mark to the
environment (Kleiman 1966, Ralls 1971,
Brown and Macdonald 1985).
Carnivores, which are mainly solitary an-
imals that patrol wide areas, primarily mark
their territories with scent (Kleiman 1966,
Gorman and Trowbridge 1989, Barja
et al. 2005). Some carnivores also use mixed
scent and visual markings. As an example
of a mixed marking, the animal scrapes the
soil, which is a well documented behaviour
in carnivores (Eaton 1970, Schaller 1972,
Bekoff 1979, Barja et al. 2005), and adds
secretions from its interdigital glands to the
visually conspicuous scrapes. Field studies
on carnivores have shown than fluctuations
in marking frequencies can indicate the on-
set of the oestrus (Wells and Bekoff 1981,
Gorman and Trowbridge 1989). Thus,
monitoring marking behaviours can indicate
reproductive physiological events and should
journal 22 v02.indb 397journal 22 v02.indb 3972010-06-18 20:20:412010-06-18 20:20:41
Isabel Barja and Francisco Javier de Miguel
be a key component of captive breeding pro-
Marking behaviour has been reported in
felines by numerous studies (Schaller 1967,
1972, Eaton 1970, Wemmer and Scow
1977, Brahmachary 1979, Brahmachary
and Dutta 1987, Smith et al. 1989, Mellem
1993, Brahmachary et al. 1999, and Brah-
machary and Singh 2000). Studies have
reported that tigers can perceive one another
through dense vegetation by sensing odours
(Schaller 1967, 1972, Gorman and Trow-
bridge 1989). Fresh marks deposited by an
animal indicate that it has recently travelled
by the marked area, and these markings can
attract or drive out visitors. For instance,
cheetahs (Acinonyx jubatus Schreber, 1975)
avoid paths that have been recently marked
by conspecifics (Eaton 1970). In some cases
the chemical composition of the marks can
provide physiological information. For exam-
ple, females in oestrus mark more frequently
than other females (Schaller 1972, Smith
et al. 1989).
Since an animal’s marking behaviour is
related to its social and physical environ-
ment, we hypothesized that closely related
species showing behavioural differences in
the wild may display different marking strat-
egies. We conducted a comparative study of
urine-marking behaviour of Siberian tigers
(Panthera tigris altaica Temminck, 1844)
and Barbary lions (Panthera leo leo Linnae-
us, 1758) at the Madrid Zoo. Siberian tigers
and Barbary lions are closely related species
with different natural habitats. To date, there
have been no comparative studies on tigers
and lions with regard to their scent marking
behaviours in the wild or in captivity. We
recorded the frequencies and the durations
of visual-scent marking behaviours to iden-
tify interspecific differences between tigers
The study was conducted on eight Sibe-
rian tigers (one adult male, two adult females,
three young males, and two young females)
and eight Barbary lions (two adult males and
six adult females) at the Madrid Zoo (Spain).
The tigers and lions occupied open enclo-
sures of 514 m2 and 730 m2, respectively.
Our observations were collected between
May 1995 and April 1996 and summed 416
hours. Marking events were recorded early
in the morning and at dawn, when the ani-
mals were most active and the probability of
marking was higher. We used continuous be-
havioural sampling (Martin and Bateson
1986) for two hours per day for two days per
week in each enclosure. We only considered
urine marking and scraping+urine squatting,
since both of these markings have recogniz-
able visual components and serve a commu-
nicative function. We also recorded the dura-
tion of each urine-marking event.
The mean urine-marking rate was signif-
icantly higher in tigers than in lions (0.5 vs
Fig. 1. Seasonal variation in the urine-marking rates of tigers and lions in captivity.
journal 22 v02.indb 398journal 22 v02.indb 3982010-06-18 20:20:412010-06-18 20:20:41
Urine marking in captive tigers and lions
0.1 marks/h, c2 = 211.8, df = 1, n = 362,
P <0.001). As shown in Figure 1, the tigers
showed the highest urine-marking rate dur-
ing the winter, followed by the spring and
autumn. The lowest urine-marking rate was
observed in the summer (c2 = 31.1, df = 3,
n = 251, P <0.001). In contrast, we did not
observe a significant seasonal variation in the
urine-marking rates of lions (Fig. 1) (c2 = 6.3,
df = 3, n = 140, P> 0.05).
We observed interspecific differences in
the duration of the urine-marking events.
The duration was shorter in tigers (1.95 ±
1.66 s, n = 88) than in lions (10.47 ± 7.82 s, n
= 21). Additionally, the variability was greater
in lions than in tigers (lions: 3.5–32 s, tigers:
1.5–2 s), and the distributions were signifi-
cantly different (Mann-Whitney U test = 58,
n = 109, P <0.001).
The results of our study demonstrate
significant differences in the urine-marking
behaviours of large felids. We observed that
urine-marking rates were significant higher
in tigers than in lions. Tigers dwell in forest
environments where scent marks, such as
urine, constitute an effective communication
channel. Tigers have been reported to have a
greater proportion of lipids in their anal se-
cretions than lions (Andersen and Vulpi-
us 1999, Burger et al. 2008), which may play
a substantial role in retaining volatiles. These
data support the hypothesis that chemical
communication is more essential for tigers
than for lions.
Tigers showed seasonal variations in their
marking rates, while lions showed no signifi-
cant variations. These differences may be due
to differences in the animals’ reproductive
cycles. Tigers in the wild show seasonal dif-
ferences in the frequency of urine markings,
and the highest marking rates occur during
the reproductive period (Schaller 1967).
According to Tilson and Seal (1987),
marking increases during oestrus and/or
proestrus. Smith et al. (1989) reported
that marking by female tigers is indicative
of oestrus, although marking ceased during
pair-bonding. Males marked more frequent-
ly when females were in oestrus. Seasonal
patterns in marking frequencies have been
documented in other felid species, where in-
creases in markings occur during the mating
season (Wielebnowsky and Brown 1998,
Mellem 1999). Therefore, we conclude that
the urine-marking rate in tigers may serve
as a good indicator of both oestrus and the
In contrast, lions do not show a seasonal
pattern in their marking frequencies in the
wild (Schaller 1972). This is likely because
female lions are annually polyoestrus, while
female tigers are seasonally polyoestrus. Ad-
ditionally, females who are annually poly-
oestrus in other carnivorous species, such as
spotted and striped hyenas, do not show sea-
sonal patterns in their marking frequencies
(Rieger 1981). In these annually polyoestrus
animals, marking behaviours are not related
to the mating time or the breeding period
The differences in the duration of urine
marking in tigers and lions may be related
to differences in their urine-marking fre-
quencies. Lions urinated with a lower fre-
quency and longer durations. Since tigers
live in tropical forests, they rely more heav-
ily upon urine marking. Logically, tigers and
other species that mark frequently should
use shorter urine-marking acts to avoid ex-
hausting their semi-chemical reserves. For
instance, Barja and Miguel (2004) re-
ported that more frequent urination events
were shorter in duration in captive wolves.
Since lions inhabit more open areas that fa-
vour visual marking, they need not urinate as
frequently. In the present study, tigers were
observed to urinate more frequently and
have short marking acts than lions. These re-
sults correspond to previous studies, which
reported that the urine-marking acts in tigers
are brief (Schaller 1967, Brahmachary
and Dutta 1987, Smith et al. 1989, Brah-
machary et al. 1991).
In conclusion, tigers and lions showed
significant differences in their urine-marking
patterns despite their exposure to similar
conditions in the zoo. One interpretation of
these results is that the conditions in the zoo
are appropriate for both species, allowing the
animals to retain the marking strategies they
used in their wild habitats.
ACKNOWLEDGMENTS: We want to ex-
press our gratitude to Madrid Zoo by their collab-
oration. We are also in debt with Cheryl Asa and
Avanti Mallapur by its useful comments.
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Isabel Barja and Francisco Javier de Miguel Download full-text
Andersen K.F., Vulpius T. 1999 – Urinary
volatile constituents of the lion, Panthera leo –
Chem. Senses. 24: 179–189.
Barja I. 2001 – La señalización en el lobo ibéri-
co (Canis lupus signatus). Comparación con
dos especies de hienas (Crocuta crocuta y Hy-
aena hyaena) – Ediciones de la Universidad
Autónoma de Madrid, Madrid, 372 pp.
Barja I., Miguel F.J. 2004 – Variation in stim-
ulus, seasonal context and response to urine
marks by captive Iberian wolves (Canis lupus
signatus) – Acta Ethol. 7: 51–57.
Barja I., Miguel F.J., Bárcena F. 2005 –
Faecal marking behaviour of Iberian wolf in
different zones of their territory – Folia Zool.
Bekoff M. 1979 – Ground scraping by male do-
mestic dogs: a composite signal – J. Mammal.
Brahmachary R.L. 1979 – The scent-marking
of tigers – Tiger Paper, 6: 19–20.
Brahmachary R.L., Datta J., Poddar-
Sarkar M. 1991 – The marking fluid of the
tiger – Mammalia, 55: 150–152.
Brahmachary R.L., Dutta J. 1987 – Chemi-
cal communication in the tiger and leopard
(In: Tigers of the world, Eds: R.L. Tilson, U.S.
Seal) – Noyes Publication, USA, pp 296–302.
Brahmachary R.L., Singh M. 2000 – Behav-
ioral and chemical aspects of scent marking in
the Asiatic lion – Curr. Sci. India, 78: 680–682.
Brahmachary R.L., Singh M., Rajput M.
1999 – Scent marking in the Asiatic lion –
Curr. Sci. India, 76: 480–481.
Brown R.E., Macdonald D.W. 1985 – Social
odours of Mammals – Clarendon Press, Ox-
ford, 386 pp.
Burger B.V., Viviers M.Z., Bekker J.P.I., le
Roux M., Fish N., Fourie W.B., Weib-
chen G. 2008 – Chemical characterization of
territorial marking fluid of male Bengal tiger,
Panthera tigris – J. Chem. Ecol. 34: 659–671.
Eaton R.L. 1970 – Group interactions, spacing
and territoriality in cheetahs – Z. Tierpsychol.
Gorman M.L., Trowbridge B.J. 1989 – The
role of odor in the social lives of carnivores
(In: Carnivore behavior, ecology and evolu-
tion, Ed: J.L. Gittleman) – Cornell University
Press, Ithaca, pp. 57–88.
Kleiman D.G. 1966 – Scent marking in the
Canidae – script Symp. Zool. Soc. Lond. 18:
Martin P., Bateson P. 1986 – Measuring be-
haviour: an introductory guide – Cambridge
University Press, Cambridge, 200 pp.
Mellem J.D. 1999 – A Comparative analysis of
scent-marking, social and reproductive be-
havior in 20 species of small cats (Felis) – Am.
Zool. 33: 151–166.
Ralls K. 1971 – Mammalian scent marking –
Science, 171: 443–449.
Rieger I. 1981 – Hyaena hyaena – Mammalian
species, 150: 1–5.
Schaller G.B. 1967 – The deer and the tiger –
University of Chicago Press, Chicago, 384 pp.
Schaller G.B. 1972 – The Serengeti lion: a
study predator-prey relations – University of
Chicago Press, Chicago, 494 pp.
Smith J.L.D., McDougal C., Miquelle D.
1989 – Scent marking in free-ranging tigers,
Panthera tigris – Anim. Behav. 37: 1–10.
Tilson R.L., Seal U.S. 1987 – Tigers of the
world – Noyes Publication, USA, 502 pp.
Wells M.C., Bekoff M. 1981 – An observa-
tional study of scent marking in coyotes, Canis
latrans – Anim. Behav. 29: 332–350.
Wemmer C., Scow K. 1977 – Communica-
tion in the Felidae with emphasis on scent
marking and contact patterns (In: How Ani-
mals Communicate, Ed: T.A. Sebeok) – Indi-
ana University Press, Bloomington (IN), pp.
Wielebnowsky N., Brown J.L. 1998 – Be-
havioral correlates of physiological estrus in
cheetahs – Zoo Biol. 17: 193–209.
Received after revision September 2009
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