Mammalian Biology 87 (2017) 36–39

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Mammalian Biology
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Short communication

Does human infrastructure shape scent marking in a solitary felid?
Miha Krofel a,∗,1 , Lan Hočevar a , Maximilian L. Allen b,1ab

Wildlife Ecology Research Group, Department of Forestry, Biotechnical Faculty, University of Ljubljana, Večna pot 83, SI-1000 Ljubljana, Slovenia
Department of Forest and Wildlife Ecology, University of Wisconsin,1630 Linden Drive, Madison, WI 53706, USAa r t i c l e

i n f oArticle history:
Received 9 November 2016
Accepted 12 May 2017
Handled by Laura Wilson
Available online 20 May 2017
Keywords:
Eurasian lynx
Human development
Lynx lynx
Roads
Urine markinga b s t r a c t
Human development, such as construction of roads and trails, can affect several animal behaviour traits
and is increasing worldwide. Despite the importance of scent marking for communication and socialorganization in many mammals, it is not clearly understood how scent marking is affected by humanhabitat modiﬁcation. We used snow tracking data collected across six winters to study the effects of roadinfrastructure on the marking behaviour of Eurasian lynx (Lynx lynx) in a human-dominated landscape.
We found that lynx marked at higher rates (11.2 urine sprays/km) on human routes (forest roads andlogging trails) than when walking in natural habitat (5.8 sprays/km). This pattern was noted in both sexes,
although males walked on human routes more frequently and scent marked more often than females.
Compared to previous studies the mean marking rates we observed were the highest documented so far
for wild felids. This may mainly be due to regular use of human routes (42% of all lynx movement) and thehigh scent marking rates found thereon. Human routes channelize lynx movement, and scent marking
on these routes likely increases the probability of the chemical message being received by a conspeciﬁc;therefore human routes might enable more effective communication in territorial felids. Since most of thelandscapes worldwide are covered by road networks and various human objects, the marking behaviourof wild mammals may be profoundly changed due to human-caused habitat modiﬁcations.
© 2017 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.

Human infrastructure, such as roads and trails, is an increasingly
widespread feature of most landscapes worldwide and can produce strong effects on animal communities (Trombulak and Frissell,
2000). These effects have been studied extensively for several animal behaviour traits, such as movement, home range size anddistribution, selection of breeding sites, and feeding behaviour (e.g.,
Ciuti et al., 2012; Jerina, 2012; Sazatornil et al., 2016; Trombulak
and Frissell, 2000). Wildlife is well known for having adverse reactions to human activity and scent (Bates et al., 2007; Wilmers
et al., 2013; Tablado and Jenni, 2015), but studies focusing on
determining how interspeciﬁc dynamics with humans, including
linear infrastructure, activity and development, affect the scentmarking behaviour of wild animals is not well understood. There
are synanthropic species that are sometimes attracted to developed areas regardless of scent, while results for species that tend
to avoid humans are mixed. For example, puma (Puma concolor)
scent-marking areas are located away from highly developed areas(Wilmers et al., 2013), while wolves (Canis lupus) regularly use
roads, but evidence of selection of this human infrastructure for∗ Corresponding author.
1E-mail addresses: miha.krofel@bf.uni-lj.si, miha.krofel@gmail.com (M. Krofel).
Contributed equally to the manuscript.

scent marking has been equivocal (Peters and Mech, 1975; Zubet al., 2003).
Scent marking is an important aspect of communication andsocial organization for many mammals, and it is the predominant
form of communication among most solitary felid species (Allenet al., 2016; Bailey, 1974; Smith et al., 1989; Vogt et al., 2014). Scent
marking has many functions, including marking territories (Bailey,
1974; Smith et al., 1989; Gosling and Roberts, 2001) and advertising
for mates (Vogt et al., 2014; Allen et al., 2015). Many species returnto speciﬁc areas located in their home ranges to scent mark; theseareas are conceptually referred to as ‘chemical bulletin boards’,
as they are repeatedly used for indirect communication throughscent marking (Ferkin and Pierce, 2007; Wolff et al., 2002). Individuals scent mark throughout their home ranges rather than just on
borders, and are likely to select scent-marking locations that areconspicuous and have a high probability of success of being foundby conspeciﬁcs (Ahlbom and Jackson, 1986; Bothma and le Riche,
1995; Gosling and Roberts, 2001; Piñeiro and Barja 2012).
To study the potential effects of road infrastructure on thescent-marking behaviour in solitary felids, we used snow tracking data collected across six winters to measure scent-marking
rates of Eurasian lynx (Lynx lynx) in a human-dominated landscape.
Eurasian lynx is a solitary apex carnivore that is found across Europe

http://dx.doi.org/10.1016/j.mambio.2017.05.003
1616-5047/© 2017 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.

M. Krofel et al. / Mammalian Biology 87 (2017) 36–39

and Asia. As is typical for solitary felids, Eurasian lynx have large
home ranges (Linnell et al., 2001), display intrasexual territoriality (Breitenmoser and Breitenmoser-Würsten, 2008; Schmidt et al.,1997), and scent mark (via urine spraying) throughout their homerange (Vogt et al., 2014). In general lynx avoid areas of intensive
human land use (Schadt et al., 2002), but regularly use forest roads
and similar habitats with less human use (Vogt et al., 2016). Lynx
sometimes scent mark more frequently in the periphery of theirhome ranges (Sæbø, 2007), and also when walking on or close totrails, roads, forest edges, ravines, and small watercourses throughout their home range (Vogt et al., 2016).
The objectives of our study were: (1) Determine whether lynx
scent mark more frequently on human routes (gravel forest roads
and dirt trails used for logging) compared to natural (off-road) habitats. We predicted higher marking frequency on human routes,
because such linear infrastructure might channelize lynx movement and thus increase their utility as chemical bulletin boards.
(2) Compare scent-marking rates between male and female lynx.
According to previous studies (Sæbø, 2007; Allen, 2014) we predicted that males would exhibit higher overall rates of scentmarking than females. (3) Perform a review of each study thatreported marking rates of felids and compare them with resultsof our study.
Our study was conducted in the forest landscape of Dinaric
Mountain Range in Slovenia (45◦ 25 –45◦ 47 N, 14◦ 15 –14◦ 50 E)
where a dense network of regularly maintained human routes isestablished (primarily for logging activities) with an average density of approximately 2 km of forest roads and 10 km of dirt trails
per km2 of forest. Eurasian lynx in Slovenia are part of the Dinaric
lynx population, one of the most threatened populations in Europe(Krofel and Jerina, 2016).
We used intensive snow tracking to document the scentmarking behaviours in Eurasian lynx. In total we followed 20 lynx
tracks, but for our analyses used the 18 tracks which were >1 km
in length (nmale = 13, nfemale = 3, nunknown = 2). These tracks were
sampled during six winters in the course of twelve years from2005 to 2017 (n05-06 = 2, n06-07 = 5, n07-08 = 3, n14-15 = 1, n15-16 = 5,
n16-17 = 2); on ﬁve occasions we followed the track across multiple days. Most (94%) of the lynx tracks alternated several times
between human routes and natural habitat. We noted the sex of thelynx when possible (n = 16) either through genetic analyses from
non-invasive samples collected on the tracks (n = 8; scats, urine
and hairs were used for genetic analysis performed on 20 polymorphic microsatellite loci and one sex-speciﬁc locus; for details see
Polanc et al., 2012 and Sindičić et al., 2013), the tracking of radiocollared individual sexed during capture (n = 1; Krofel et al., 2013),
or the characteristics of tracks (n = 7; size of the foot print and presence/absence of kitten tracks). Because it was not possible to always
individually identify the lynx (n = 10), we could not determine thenumber of different individuals sampled and the same individuals
were likely tracked multiple times during the study period.
We focused on urine spraying, because cheek rubbing was difﬁcult to detect, Eurasian lynx generally do not use scrapes or faeces
for marking (Breitenmoser and Breitenmoser-Würsten, 2008), andclaw marking was infrequent (n = 2). When urine marking, lynx typically create an easily recognizable track pattern in the snow bymaking a short detour from their direction of travel and turning
their hindquarters towards an object in order to spray urine uponit (Vogt et al., 2016; see also Appendix A). After new snowfall welocated a fresh lynx track and followed it while documenting thelocation of each occurrence of urine spraying. We used a handheld
GPS to record the course and length of the lynx path, and notedthe coordinates for each instance of urine spraying. We also documented whether the sprayed urine occurred on a human route (i.e.,
gravel road or logging dirt trail) or in natural habitat (i.e. off-road).

37

Fig. 1. The mean scent-marking rates of Eurasian lynx, split by all lynx, males, andfemales, and shown among natural habitats (white), human routes (black), and mean
for all areas (grey), with error bars representing the standard error.

We used program R version 3.2.2 (R Core Team 2015) for all of
our statistical analyses, and in each analysis we considered p < 0.05
to be statistically signiﬁcant. We ﬁrst calculated summary statistics
for our data, calculated marking rates as number of urine sprays/kmof lynx track, and then tested our predictions. We calculated scentmarking rates by dividing the number of recorded urine sprays by
the length of lynx path for each track and separately for each sex
and habitat type. We used non-parametric tests due to small sample sizes. We ﬁrst used a Wilcoxon Signed Rank Test to determine
whether lynx differ in their rate of urine spraying between humanroutes and natural habitats. Second, we used a Mann-Whitney U
Test to determine whether male and female lynx differ in their rateof urine spraying. Third, we used Wilcoxon Signed Rank Tests todetermine whether either male or female lynx differ in their rateof urine spraying between human routes and natural habitats.
We followed the 18 Eurasian lynx tracks a total of 94.1 km andrecorded 619 urine marking events. Lynx walked 54.2 km in natural
habitat (58%), and 39.9 km on human routes (42%; 30.2 km on gravel
forest roads and 9.7 km on logging dirt trails). We followed malelynx for a total of 61.6 km, with 27.1 km walking in natural habitat(46%) and 33.4 km on human routes (54%). We followed female lynx
a total of 27.6 km, with 23.3 km walking in natural habitat (85%) and4.2 km on human routes (15%).
Overall, we documented a mean urine marking rate of 7.1(±1.4 SE, range: 0.0–20.0) sprays/km. We found that lynx marked
with higher rate on human routes (11.2 sprays/km, ±2.1 SE) thanwhen walking in natural habitat (5.8 sprays/km, ±1.6 SE) (V = 116,
p = 0.014) (Fig. 1). We also found that males sprayed urine signiﬁcantly more frequently (9.1 sprays/km, ± 1.6 SE) than females (1.1
sprays/km, ±1.0 SE) (W = 1, p = 0.007) (Fig. 1). Both sexes exhibited an increased frequency of scent marking on human routes(male = 14.0 sprays/km, ±2.5 SE; female = 3.6 sprays/km, ±3.6 SE)
when compared to natural habitats (male = 7.4 sprays/km, ±1.9 SE;
female = 0.5 sprays/km, ±0.5 SE) (Fig. 1), but only males were signiﬁcantly different between types (V = 77, p = 0.027).
Compared to previous studies of scent-marking rates of felids,
the overall mean urine spraying rates for Eurasian lynx in Slovenia
were the highest marking rates reported so far, with the exceptionof studies that only reported the maximum rates found (Table 1).
The scent-marking rates we recorded in natural habitats were similar to previous studies of lynx and other felids, but marking rates werecorded on human routes were over 3-times as high as other documented marking rates, and resulted in our overall lynx marking
rates being twice as high as any other previous study. Our review
of reported scent-marking rates for felids also highlights the needThis Study
This Study
This StudySæbø (2007)
Vogt et al. (2016)
Hücht-Ciorga (1988)
This Study
Saunders, 1963
Bailey (1974)
Bothma and le Riche (1995)
Bothma and le Riche (1995)
Bothma and le Riche (1995)
Schaller (1977)
Smith et al. (1989)
Schaller (1972)
Molteno et al. (1998)
n.a./94 km
n.a./94 km
n.a./94 km
n.a./ca. 800 km
15/185 km
2/105 kmn.a./94 km
n.a./n.a.
2/2 kmn.a./2665 km
n.a./2665 km
n.a./2665 km
2/10 km2/303 km
1/2 km
1/1 kmWinter
Winter
Winter
Winter
Winter
Winter
Winter
Wintern.a.
All yearAll yearAll yearn.a.
All yearn.a.
June/JulyAll
Natural Habitats
Human Routes
All
All
All
All
All
All
All
All
All
All
Animal trail
Road
All
Urine Spraying
Urine Spraying
Urine Spraying
Urine Spraying
Urine Spraying
Urine Spraying
Claw Marking
Urine Spraying
Urine Spraying
Claw Marking
Urine Spraying
Body Rubbing
Scraping
Urine Spraying
Urine Spraying
Urine Spraying
1.10
0.52
3.59
0.94
n.a.
n.a.
n.a.
n.a.
n.a.
0.09
0.01
0.04
n.a.
1.80
n.a.
64.6a
a

only maximum rates reported.
only 2 observations.
2
only 1 observation.

9.09
7.38
14.00
1.81
n.a.
n.a.
n.a.
n.a.
n.a.
0.13
0.10
0.07
0.301
2.40
3.682
n.a.
1

All

7.10
5.83
11.19
1.35
3.40
1.03
0.02
10.6–11.8a
3.001
0.12
0.08
0.06
n.a.
2.11
n.a.
n.a.
Lynx lynx
Lynx lynx
Lynx lynx
Lynx lynx
Lynx lynx
Lynx lynx
Lynx lynx
Lynx canadensis
Lynx rufus
Panthera pardus
Panthera pardus
Panthera pardus
Panthera uncia
Panthera tigris
Panthera leo
Felis nigripes

Species

Rate (scent marks/km)

Male

FemaleType of Marking

SubsetSeasonSample size (number ofanimals/total track length)

SourceM. Krofel et al. / Mammalian Biology 87 (2017) 36–39
Table 1The scent-marking rates (scent marks/km) for felids noted in the scientiﬁc literature. When possible, for each study we provide the rates for all individuals, males, and females, the rates for each type of scent marking, and noteson where and when they occurred and the samples sizes.

38

for future studies that focus on marking rates in different types ofhabitat and among types of scent marking other than urine spraying(Table 1).
Since many of the previous studies of lynx and of other felid
species were conducted in regions with lower road density compared to our study area, we suggest that the intensity of humanhabitat modiﬁcation could be a factor in anthropogenic perturbations of the scent-marking behaviour of solitary felids. At the sametime, we assume that the low intensity of human use of this infrastructure plays an important role, as felids are known to avoid areas
with frequent human presence (Schadt et al., 2002; Wilmers et al.,2013). Therefore the combination of a dense network of humaninfrastructure that has low human use might present the situation
with pronounced effects on felid marking behaviour.
The high overall marking rates we observed are primarily due
to male lynx, which more frequently walked on human routes andalso scent marked more frequently than females. This difference
among sexes is similar to other felids (Allen et al., 2015; Bothma
and le Riche, 1995), and supports similar ﬁndings for Eurasian lynx
where males are more frequent visitors to scent-marking sites, andalso scent mark more frequently than females (Vogt et al., 2014,2016). Human routes do not delineate the home range boundaries
recorded in the study area (Krofel, 2012), and lynx are using humanroutes throughout their home ranges, likely for ease of movement
and to patrol their territories. Therefore, the more frequent use ofhuman routes by males would be expected given their larger homerange size compared to females (Breitenmoser and BreitenmoserWürsten, 2008; Linnell et al., 2001). Scent marking by lynx shows
seasonal variation (Schmidt and Kowalczyk, 2006; Vogt et al., 2014,2016), with a peak during the late winter breeding season and theleast frequent marking in summer when mothers are denning andraising young (Vogt et al., 2014). Because this study was conductedduring winter, some of our tracking days were likely to correspondwith the breeding season and the higher scent-marking rates that
occur for both sexes during this period could be a confounding
factor when comparing with other studies.
Besides providing a means of easier and faster travel, the linear human infrastructure could also increase the efﬁciency of scentmarking. A recent study from Switzerland showed that lynx scentmark more intensively when walking in a straight line (Vogt et al.,2014). In a landscape with rugged terrain, which is characteristic for most of the range of the Dinaric lynx population, thepossibility of walking in a straight line is notably more likely onhuman routes than in natural habitat. Additionally, the frequent
use of human routes channelizes movement, and may provide
structure for ‘chemical bulletin boards’ used by the mammal populations throughout their home ranges (e.g., Ferkin and Pierce,
2007; Vogt et al., 2014; Wolff et al., 2002). Scent marking on
human routes therefore likely increases the probability of a chemical message being received by a conspeciﬁc, and we suggest thathuman modiﬁcation of the landscape may enable territorial felids
to communicate more effectively. Given the fact that most of thelandscapes in the modern world are now covered by road networks (Trombulak and Frissell, 2000) the scent-marking behaviourof species such as lynx that selectively use human routes for indirect communication may have been profoundly changed due tohuman-caused habitat modiﬁcation.
Besides creating linear infrastructure, humans could also affectfelid marking behaviour through the construction of cabins, fences,
poles, and other conspicuous objects. Like many other carnivores,
lynx often use conspicuous objects in their environment for scentmarking, and human objects often stand out in natural habitats andlikely serve as attractive objects for scent marking (for example
see video ﬁle showing Eurasian lynx scent marking forest cabin in
Appendix A). Indeed, a study in Poland revealed that human objects
were the most frequently used objects by Eurasian lynx for scentM. Krofel et al. / Mammalian Biology 87 (2017) 36–39

marking by cheek rubbing (Schmidt and Kowalczyk, 2006). Sincesuch objects are often located along human routes, linear infrastructure likely promotes their use as marking objects.
Among solitary felids, scent marking is essential to territoriality(Bailey, 1974; Gosling and Roberts, 2001; Smith et al., 1989) andmate selection (Allen et al., 2015; Vogt et al., 2014). Our ﬁndings
suggest that Eurasian lynx possibly adapted their scent-marking
behaviour to human co-existence over the last thousands of years
as human infrastructure has developed and increased across theEurasian landscape. This highlights the need for more studies tobetter understand anthropogenic effects on scent marking of carnivores in human-dominated landscapes. Roads and other linear
human infrastructure can have strong negative effects on carnivores, including collisions with vehicles, habitat fragmentation,
limiting dispersal and gene ﬂow, as well as increasing the accessof people to remote areas which might increase legal and illegal
hunting (see Trombulak and Frissell, 2000 for review). Our results
suggest that human routes with low use by people might improve
effectiveness of indirect communication in solitary felids. However,it remains to be seen whether this could have beneﬁcial impacts onimportant traits at the population level, such as increased breeding
success and stability of the social system, and whether these effects
are large enough to counteract the negative effects of human routes
and other development. Studies to determine scent-marking rates
are labour-intensive, but can provide information on the function
and mechanisms of scent marking, and we encourage future studies of this kind, especially of species and marking behaviours forwhich marking rates are poorly known.
Acknowledgements
This work was partly funded by the Slovenian Research Agency(grant no. P4-0059) and by the European Union through INTERREG
IIIA Neighborhood Program Slovenia/Hungary/Croatia 2004–2006(project “DinaRis”). We are grateful to the three anonymous
reviewers for reviewing early drafts and providing several usefulsuggestions that improved the manuscript.
Appendix A. Supplementary data
Supplementary data associated with this article can be found, inthe online version, at http://dx.doi.org/10.1016/j.mambio.2017.05.
003.
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