SCENT MARKING IN FALLOW DEER: EFFECTS OF LEKKING
BEHAVIOR ON RUBBING AND WALLOWING
GIOVANNA MASSE! AND
R.
TERRY BOWYER
Institute of Terrestrial Ecology, Hill of Brathens, Banchory, AB31 4BY, Scotland.
United Kingdom (GM)
Institute of Arctic Biology, and Department of Biology and Wildlife, University of Alaska Fairbanks,
Fairbanks, AK, 99775 (RTB)
Present address of GM: Central Sciences Laboratory, Sand Hutton, York, Y04 lLZ
United Kingdom
We studied scent marking (rubbing of trees and wallowing) by lekking fallow deer (Dama
dama) in central Italy during October 1995. We hypothesized that composition of tree
species and their location and physical properties affected scent marking and thereby played
a role in the selection of sites for leks. Wallows of fallow deer were located adjacent to
rubbed trees; wallows in 73% of 200 sites occurred within ::::::::3 m from a tree. Deer wallowed
preferentially near Pistacia and Juniperus. Trees rubbed by deer occurred more often along
roads (24%) than inside the woodland (5.8%). Only Pistacia was rubbed more often that
its proportional occurrence in the environment. Regardless of tree species, fallow deer
selected those trees- with the greatest height, largest diameter, and greatest number of stems
for scent marking. The species composition of trees on two leks differed from surrounding
areas. Fallow deer rubbed trees differently inside and outside of leks, and within leks they
selected Juniperus and Myrtu~. The close spatial association of wallows and rubbed trees,
however, indicated the function of these two scent marks may be related.
Key words: Dama dama, fallow deer, scent marking, Jekking, wallowing, rubbing, rutting,
Cervidae, Italy
yer et al., 1994). Large dominant males
tend to scent mark more frequently than
smaller subordinates (Bowyer, 1986; Bowyer and Kitchen, 1987; Miquelle, 1991;
Nielsen et aI., 1982).
In southern Europe, fallow deer (Dama
dama) congregate in traditional areas (leks)
where males exhibit specialized territorial
behavior at the end of September (Apollonio et al., 1992; Balmford et al., 1993).
During that period, adult males debark trees
with their antlers and rub their foreheads
and preorbital glands on those trees. These
males also spray urine on their bellies,
scrape the ground with front legs and wallow (Chapman and Chapman, 1975; Walther, 1984). Wallows are small depressions
made by males in the ground where they
may urinate and lie during rut. Fallow deer
Scent marking is common among both
territorial ungulates and nonterritorial mammals (Gilbert, 1973; Gosling, 1987; Ralls,
1971). Among ungulates, visual (rubs,
scrapes, and wallows), olfactory (urine and
pheromones), and acoustic (vocalizations)
signals are used to communicate with conspecifics (Bowyer et aI., 1994; Coblentz,
1976). Visual and olfactory cues often are
located on conspicuous landmarks, such as
edges of forests, along roads, and at elevated sites. Scent marks may be important
in conveying information on body size and
strength of males, or in priming estrus of
females (Bowyer and Kitchen, 1987;
McCullough, 1969; Miquelle, 1991; Sawyer et aI., 1989). Many of these scent-marking behaviors are linked or sequenced differently among species of Cervidae (Bowjoum,,{ of M{mmU/I(J~y. 80{2):633--638. 1999
633
634
JOURNAL OF MAMMALOGY
possess several glands that may aid in scent
marking (Chapman and Chapman, 1975;
Kennaugh et a!., 1977).
Although many studies have investigated
the lekking behavior of fallow deer in terms
of their reproductive strategy, social groupings. and inter- and intra-sexual interactions
(Alvarez et a1.. 1990; Buschhaus et aI.,
1990; Pemberton and Balmford, 1987), no
information is available on the physical
characteristics or spatial arrangement of
scent marks, or on factors affecting selection of trees for scent marking.
We analyzed the scent marks of fallow
deer to test the following hypotheses: 1)
trees along roads were marked with greater
frequency than trees within the forest; 2)
deer rubbed (scent marked) the largest and
most aromatic trees available; 3) wallows
were spatially associated with rubbed trees;
and 4) trees in leks were marked in greater
proportion than trees outside leks.
MATERIALS AND METHODS
Study area.~This study was conducted in the
Maremma Natural Park along the coast of central
Italy (42°39'N, 11°05'E). Precipitation ranged between 500-750 mm/year. Mean temperature was
24°e in July and 6°e in February. The area was
dominated by Mediterranean maquis, characterized by Quercus ilex, Arbutus unedo, Pistada
lentiscus. and Rosmarinus officinalis. We collected data in pinewood habitat, which encompassed
ca. 600 ha, and was characterized by Pinus pinea,
Pistacia lentiscus, Phillyrea angustifolia, and
Myrtus communis. A detailed description of the
vegetation was provided by Arrigoni et a1.
(1976). Roads, covering <5% of the study area,
were rarely used by vehicles.
Sampling procedure.-Data were collected
during the first 2 weeks of October 1995, when
fallow deer were rutting. We located tree species
every 2 m along transects to sample their availability and use (rubbing) for scent marking.
Fourteen transects, each 50 m in length, were
used by randomly positioning a 50 m rope. Transects were either 2:50 m into pinewood (n = 9)
or along roads (n = 5). For each tree or shrub
that hit the rope at 2-m intervals, we recorded
species, maximum height (em), circumference
(cm) at 20 cm above the ground, and number of
Vol. 80, No.2
stems at a point 1.3 m above the ground (or the
terminal number of stems for plants < 1.3 m in
height). We chose two leks, recogni~able from
the high concentration of adult males and their
characteristic territorial behavior on those sites
during rut. We also selected two adjacent areas
(> 150 m from the leks) where neither lelling
nor rut-related vocalizations had occurred in the
previous 4 years. Leks were generally circular
with a diameter of ca. 80-120 m, and the two
we sampled were located ca. 2 km apart.
In October 1995, 2:12 adult males were present together on one lek. Males were not counted
on the second lek, although animals were observed and heard vocalizing regularly on that
rutting ground. In and around the first lek, we
systematically searched for wallows. We collected the following data for each wallow: nearest distance to the first rubbed tree; the species
of that tree; maximum height and circumference
of the rub; maximum height of any broken
branches; and percentage of branches damaged
(rubbed or broken). We sampled rubbing (use)
and availability of trees inside and outside of the
two leks on 13 additional transects, each 75 m
in length, that we randomly located (n = 6 on
leks, n = 7 outside of leks).
Statistical analyses.-We used a two-sample
Z-test for proportions (Zar, 1984), or a chisquare test to make comparisons between trees
scent marked by fallow deer and their relative
availability in the environment (Remington and
Schork, 1970). We corrected for multiple comparisons using a sequential Bonferroni procedure (Rice, 1989), Similarly, we used at-test
(Zar, 1984) for comparing morphological characteristics of trees that deer rubbed with trees
that they did not mark. Categories of increasing
distance between wallows and rubbed trees were
evaluated with a chi-square test (Zar, 1984). We
used a multi-dimensional chi-square to test for
differences in selection of trees for rubbing by
fallow deer inside and outside of lekking areas
(Zar, 1984).
RESULTS
Wallows of fallow deer (n ~ 200) occurred within 15 m of the nearest rubbed
tree; 73% of those scent marks was within
3 m of such trees. Spatial distribution of
wallows with respect to trees differed from
expected (Fig. 1). Deer wallowed prefer-
MASSEr AND BOWYER-SCENT MARKING BY FALLOW DEER
May 1999
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,.3-4
>4-5
,.5-6
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DISTANCE BETWEEN WALLOWS AND NEAREST RUBBED TREE (m)
FIG. I.-Nearest distance between wallows of
fallow deer and rubbed trees, October 1995, Parco Naturale della Maremma, Italy.
entially near Pistacia and Juniperus but
avoided (use was less than availability) Erica and Rosmarinus (Fig. 2)_
Rubbing of trees occurred more often
along roads (24% of 125 trees) than inside
(> 50 m) the woodland (5.8% of 225 trees;
Z = 4.32, P < 0.001). Among those trees
rubbed, only Pistacia was used more than
its proportional availability (Fig. 3). Deer
scent marked trees (Table 1) with the greatest height (t = 4.394, dj = 346, P <
0.001), the largest diameter (t = 4.075, dj
= 346, P < 0.00l), and the greater number
of stems (t = 2.568, dj = 346, P < 0.05).
That pattern of scent marking was common
to species of trees rubbed by deer, except
for Rosmarinus, which was the shortest
sluub, and Juniperus, which was the tallest
species rubbed by deer (Table 1). Mean
maximum height (±SD) above the ground
of the area debarked during rubbing was
118.8 ± 41.1 cm, maximum height of broken branches was 111.1 ± 42.6 cm, and
mean diameter of broken branches was 0.6
± 0.38 em. A mean of 43.2 ± 31.7% of
branches on rubbed trees was damaged
(broken or debarked) by deer.
Composition of tree species growing on
leks differed from surrounding areas (Table
2), and fallow deer rubbed a greater proportion of trees inside (17% of 262 trees)
than outside (5% of 321 trees) of leks (Z =
3.38, P < 0.0001). Consequently, we tested
effects of lekking behavior on scent mark-
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FrG. 2.-a) Trees adjacent to wallows of fallow deer in relation to availability of trees and
b) trees available and rubbed by deer. October
1995, Parco Naturale della Maremma, Italy. An
asterisk indicates P s; 0.05 following a Bonferroni correction of a partial chi-square test. Pinus
was omitted from these analyses because of its
small sample.
z
o
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INSIDE LEK (!l- 262)
~
OUT$lDE LI'.K (!l- 321)
X 2 ~ 71.82, d.t.
P< 0.001
g
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GENERA OF TREES
FIG. 3.-Trees selected (used minus available)
by fallow deer for rubbing inside and outside of
leks, October 1995, Parco Naturale della Maremma, Italy. A partial chi-square test was performed following a significant (P < 0.001) overall test. Pinus and Rosmarinus were omitted
from this analysis to meet assumptions of the
chi-square test.
JOURNAL OF MAMMALOGY
636
Vol. 80, No.2
TABLE l.~Morphological characteristics of trees rubbed by fallow deer compared with trees that
were not rubbed. October 1995, Parco Naturale della Maremma, Italy.
Rubbed
Not rubbed
Number
Height
Species
n
Phillyrea angustifolia
Erica multiflora
4
11
2
10
26
4
Rosmarinus officina/is
Myrtus communis
Pistada lentisellS
Juniperus oxycedrus
All species
X
SD
202.5 65.5
144.5 65.2
40.0 14.1
183.0 83.4
184.2 107.6
185.0 176.9
57 174.8 100.0
Diameter
X SD
4.4
1.4
1.6
3.0
4.8
4.5
3.7
1.8
0.6
0.9
2.1
4.1·
6.6
3.6
ing by fallow deer by first examining the
mutual independence of species of trees,
whether these trees were rubbed or not, and
whether trees were located inside or outside
of the leks. The overall test was highly significant (X' ~ 71.24, d.! ~ 13, P < 0.001).
We further determined the relationship
among those variables with a test of partial
independence; fallow deer selected trees to
rub differently inside and outside of leks
(Fig. 3).
DISCUSSION
Scent marking is an important component in the behavioral repertoire of adult
male fallow deer; these cervids spent 22%
of their active time during rut engaged in
this behavior (Braza et aI., 1986). Our data
on selection of trees for rubbing and locaTABLE 2.-Percentage of trees inside and outside of two lekking areas of fallow deer, Parco
Naturale della Maremma, Italy, October 1995.
Species composition differed significantly inside
and outside of leks (X 2 = 9.96, d.f. = 4, P <
0.05).
Species of trees
(n = 255)
Inside lek
Outside lek
(n = 312)
Phillyrea angu.wifolia
Erica multiflora
Myrtus communis
Pistacia ienti.fcuoS
Juniperus oxycedrus
5.9
9.4
26.7
54.9
3.1
5.4
18.6
26.0
7.1
2.9
of stems
X
SD
Height
n
X
SD
10.5 8.3 34 161.9 154.6
14.8 9.9 81 103.0 49.9
6.5
0.7 25 62.0 25.8
14.7 6.1
76
13.7 66.9
24.7 12.1 68 127.2 78.8
7 245.7 143.0
7.5 8.5
17.9 13.3 291 118.4 86.2
Diameter
X SD
3.7
0.8
1.0
1.8
2.9
6.9
2.1
Number
of stems
X
SD
4.4
0.6
0.6
1.7
2.5
4.2
9.9 13.4
8.9
7.7 4.9
14.6 12.3
17.0 16.7
12.3 5.8
2.5
13.2
12.1
12.5
tion of wallows adjacent to such trees (Fig.
2), together with the spatial association of
those scent marks (Fig. 1), indicate their
relevance in olfactory and perhaps visual
communication to fallow deer during rut.
Likewise, males on leks selecting trees to
scent mark differently than males outside
leks (Fig. 3 and Table 2) further highlights
the importance of rubbing in the mating
system of these deer. Indeed. habitat structure is thought to affect the social organization and behavior of ungulates (Hirth,
1977; Jarman, 1974; Molvar and Bowyer,
1994). We hypothesize that the composition
of tree species on leks of fallow deer played
a role in deer selecting those area for rutting
grounds. In our study, fallow deer showed
a tendency to rub trees near roads; this
might be explained by trees on the road
edge being more accessible than those inside the dense vegetation of the pinewood.
Rubs along roads were highly visible and
probably more effective as olfactory clues
than rubs inside the wood.
We observed that rubbing of trees was
concentrated in kkking areas. Similarly,
Schaal (1986) noted that scent marks were
concentrated on leks. Chapman and Chapman (1975). however, reported that there
was no relationship between distribution of
rubbed trees and occurrence of rutting areas. This difference between studies likely
is explained by male fallow deer exhibiting
May 1999
MASSEI AND BOWYER-SCENT MARKING BY FALLOW DEER
alternative mating strategies that include
lekking, males defending large, single territories, or males being nonterritorial.
Adopting a particular strategy for mating
may depend on population density, sex ratio, and habitat structure (Apollonio et aI.,
1992; elutton-Brock et aI., 1988). In our
study area, population density of fallow
deer was relatively high (28.5 individuals!
km 2-p' V. Genov and G. Massei, in litt.)
and similar to the density of 25 fallow deer/
km' reported by Apollonio (1989) for a
population that also exhibited lekking behavior.
Our data suggest that deer selected trees
for scent marking in relation to several interrelated factors: tree location, tree species,
tree morphology and aromatic characteristics. Deer rubbed more trees along roads
than inside the woodland, and more trees
inside than outside of leks. On leks, deer
selected those species rich in essential oils
such as Myrtus and Juniperus, whereas outside of leks Pistacia was the preferred species for scent marking. Erica and Phillyrea
were avoided both on leks and in surrounding areas (Fig. 3). Similarly, deer wallowed
close to Pistacia and Juniperus but avoided
Erica, Myrtus, and Rosmarinus. Deer preferentially marked large, tall trees, and this
may explain why low-growing Rosmarinus,
which is rich in essential oils, was avoided.
Morphology of the tree was probably more
important than its aromatic characteristics
in detennining which species were scent
marked. Johansson-et al. (1995) also noted
that male roe deer (Capreolus capreolus)
selected a combination of tree characteristics such as large stems and aromatic properties for rubbing. Likewise, they reported
a preference for Juniperus. This pattern of
selection for both size and aromatic properties of trees for rubbing may explain variable results obtained for white-tailed deer
(Odocoileus virginianus) relative to which
species of trees were scent marked (Benner
and Bowyer, 1988; Kile and Marchinton,
1977; Oehler et aI., (995). The concentration of ::S;30 adult male fallow deer on a lek
637
of a few hectares (Schaal, (986), and the
amount of time they spend scent marking
during the rut (Braza et a1. 1986), may explain the high density of rubbed trees within
leks.
Whatever the functional significance of
wallowing and rubbing of trees, the close
spatial association of these scent marks on
our study areas suggests that they have a
similar function for fallow deer. Among
North American elk (Cervus elaphus), wallowing and rubbing of trees are not linked
(Bowyer and Kitchen, 1987). American bison (Bison bison) however, rub trees in
close proximity to wallows, further suggesting that these scent-marking behaviors
are linked (Bowyer et al., (998). The degree to which the linking of these behaviors
occurs in other ungulates is uncertain and
needs additional investigation.
ACKNOWLEDGMENTS
This research was funded in part by the Institute of Terrestrial Ecology, Banchory, Scotland,
and the Institute of Arctic Biology at the University of Alaska Fairbanks. We thank D. K.
Person for assistance with statistical analyses.
We are grateful to the comments of P. Beier and
an anonymous reviewer that greatly improved
the manuscript.
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Submitted 20 October 1997. Accepted 28 June 1998.
Associate Editor was John A. UtvaUis.