Forest Ecology and Management 255 (2008) 468–475

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Winter habitat use by red and roe deer in pine-dominated forest

J. Borkowski a,*, J. Ukalska b,1
a
Department of Forest Ecology and Wildlife Management, Forest Research Institute, Se˛kocin-Las, 05-090 Raszyn, Poland
b
Department of Biometry, Faculty of Agriculture and Biology, Warsaw University of Life Sciences, Nowoursynowska 159, 02-776 Warsaw, Poland
Received 31 October 2006; received in revised form 23 May 2007; accepted 5 September 2007

Abstract
Although pines (Pinus spp.) represent the main tree genus in Europe, most studies of habitat use by European deer (Cervidae) have been
conducted in spruce-dominated forests. This neglects the fact that the two forest types are quite different. Ground vegetation is much more
abundant in pine stands, especially when they are mature. In this study, red and roe deer pellet groups were counted in approximately 12,500 ha of
pine forests during the early spring over 4 years. Combinations of food and cover determined habitat attractiveness. Those habitat types which
offered both food and cover were used most intensely. The role of forage was important only where cover was sufficient, suggesting that cover plays
a primary role in influencing winter habitat use by deer.
Habitat use by red and roe deer was similar both among habitats and within them, but the smaller species, roe deer seemed able to satisfy their
cover requirements more easily than the larger red deer. No evidence of interspecific influence of red on roe deer (as suggested by Latham et al.,
1997) was found in this study. As predicted, mature pine stands were an attractive habitat for deer. It was concluded that, in hunted deer populations,
the presence of cover is important even in areas lacking large predators. Introduction of forest understories into mature pine forests should thus be
promoted in big game management.
# 2007 Elsevier B.V. All rights reserved.

Keywords: Central Europe; Cover; Winter; Habitat use; Pine forest; Poland; Red deer; Roe deer

1. Introduction ground vegetation in pine stands is generally much richer than

Red deer (Cervus elaphus) and roe deer (Capreolus
capreolus) are the most numerous and widespread deer in
Europe, and are the most extensively studied ungulates (Clutton-
Brock et al., 1982; Andersen et al., 1998). There are many studies
of habitat use by these two species in European forests (e.g.
Staines and Welch, 1984; Catt and Staines, 1987; Thirgood and
Staines, 1989; Aulak and Babińska-Werka, 1990a; Welch et al.,
1990; Latham et al., 1996; Tufto et al., 1996; Palmer and
Truscott, 2003). However, most studies have been conducted in
spruce-dominated (Picea) forests, even though the most
abundant trees in Europe are pines (Pinus spp.; UN-ECE/
FAO, 1992). There are distinct differences between pine and
spruce, especially in mature forests. Due to different crown
structure and light conditions (Robakowski et al., 2004), the
* Corresponding author. Tel.: +48 22 7150412; fax: +48 22 7150417.
E-mail addresses: boku@ibles.waw.pl (J. Borkowski),
joanna.ukalska@agrobiol.sggw.waw.pl (J. Ukalska).
1
Tel.: +48 22 5932730; fax: +48 22 5932722.
0378-1127/$ – see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2007.09.013
under the spruce canopy. Since old stands are among the most
food-abundant age classes in pine-dominated forests (Bor-
kowski, 2003), the question arises as to how deer use old pine
stands in comparison with younger stands. Old growth pine
forests are expected to be more attractive to deer than their spruce
counterparts. Moreover, most studies on red and roe deer habitat
use come from Scotland. Only a few were conducted in winter,
which is a period of physiological stress, affecting important
processes such as population dynamics (among others). Winter is
also the season damage caused by deer in forests is most
concentrated (Gill, 1992). Thus, a better understanding of factors
affecting winter habitat use by deer is important for both game
and forest management.
In general, habitat use by deer is determined by the presence
of both food and cover. The role of cover in habitat use may be
especially important in winter, when cervids reduce their food
intake and live to a remarkable extent off fat reserves (Putman,
1988). Two recognized cover types are thermal cover, in which
a forest overstorey protects against weather and sun, and hiding
(security) cover used in escape and as protection against
predators and humans (Peek et al., 1982). Although many
 J. Borkowski, J. Ukalska / Forest Ecology and Management 255 (2008) 468–475
studies have documented the role of thermal cover (see
Mysterud and Østbye, 1999, for references), recent experi-
mental study demonstrated no positive effect of thermal cover
on elk (C. elaphus nelsoni; Cook et al., 1998). On the contrary,
the authors found that thermal cover resulted in greater over
winter mass loss, fat catabolism and mortality. Less information
is available on hiding cover, which is generally presumed to be
associated with predation risk (Mysterud and Østbye, 1999). In
fact, it is often people who influence deer behaviour and habitat
use most. Hunting, as the extreme manifestation of human
influence, is obviously an important factor. While animals in
unhunted populations are usually tolerant of human presence
(Schultz and Bailey, 1978; Borkowski, 2001), hunting tends to
promote flight behaviour (Altman, 1958; Behrend and Lubeck,
1968). Moreover, as hunting is ‘‘human predation,’’ it ensures
that hiding cover will be an important determinant of forest
habitat use by deer, even where natural enemies are absent. This
has been demonstrated for red deer by Bonenfant et al. (2004).
Kufeld et al. (1988), found that mule deer (Odocoileus
hemionus) used habitats with increasingly better cover as the
hunting season progressed. A reasonable assumption, is that use
of old pine forest (a relatively open habitat) during the hunting
season would be positively influenced by understorey cover
conditions.
To date, the role of cover in habitat use among European
deer, has mostly been studied in roe deer (Mysterud and
Østbye, 1995), with much less attention given to red deer.
Being much larger than roe deer, red deer may experience
greater difficulty finding cover under the same habitat
conditions. For instance, habitat use by red deer in young
pine stands depends much more on tree height than is the case
with roe deer (Borkowski, 2004). Similar phenomena may also
apply in older forests.
In spite of the numerous studies on both deer species,
relatively little information is available on comparisons of
habitat use and potential competition. However, Latham et al.
(1996) compared red and roe deer densities in 20 localities and
found a negative correlation between them. In the same 20
forests, the authors also found that red deer density had a
negative influence on roe deer density, while the reverse was not
the case (Latham et al., 1997). Both these studies are from
Scotland and there is little information available from other
places. If there is an interspecific influence of red on roe deer,
spatial distribution of both species may be negatively related,
not only between areas but also within them.
The objective of this study was to examine patterns of winter
habitat use by red and roe deer in pine-dominated forests. It was
predicted that in pine-dominated forests, deer would use mature
stands more frequently as compared to deer in spruce forests.
Another prediction was that a hunted population would use
habitats with security cover more intensely than habitats with
less security cover, even in predator-free environments. It was
also predicted that security cover would influence habitat use
more in large deer species (red deer) than in small ones (roe
deer). The final prediction was that, if interspecific competition
between them is a factor, than deer densities in different
habitats within one area would be negatively correlated.
469
2. Study area
The study was carried out near Rudy Raciborskie, in a Forest
District of ca. 17,500 ha in southwestern Poland. As a large
stand-replacement fire burned about 5000 ha of the District in
1992, data referred to here are from unburned forest only.
Coniferous habitat types accounted for approximately 65% of
the study area, with Scots pine (Pinus silvestris), the main tree
species in the canopy (up to 85% of cover). As there was almost
no management in the unburned forest for several years after
the fire, the proportion of new plantations, and especially of
pre-thickets, was minor. All of the forest is managed for timber
with artificial regeneration accounting for the greatest part of it.
Brown podzolic soils, together with podzols, make up 73% of
the soils in the study area. About 20 years ago, foresters in this
area began to introduce understories as part of a nationwide
strategy to enrich homogenous pine stands. The main species
used for this purpose were Norway spruce (Picea abies),
common beech (Fagus sylvatica), black dogwood (Frangula
alnus) and black cherry (Prunus serotina).
The winter climate of the study area is very mild. The
average temperature between December and March is 0.2 8C,
with January being the coldest month (2.1 8C). The number of
days with snow covers between December and February is
usually around 45, and average snow depth rarely exceeds
5 cm. The area is flat with an elevation range between 180 and
307 m above sea level.
At the time of the study, red and roe deer population
densities (as estimated by drive counts) were of 4–6 and 6–10
individuals/100 ha of forest area, respectively (P. Nasiadka,
unpubl. data). Other ungulates in the area were wild boar (Sus
scrofa) and the less-common fallow deer (Dama dama). All the
species were hunted by both individuals and groups. The
hunting season varied with the species, but was most intense
between October and February. There are no large predators in
the area, except for occasional occurrences of single wolves
(Canis lupus). The study area is located in Silesia, a densely
populated, industrial region of Poland, and is commonly used
for recreation.
3. Methods
The study was conducted between 1997 and 2000, using
standing crop counts of faecal pellet groups (Neff, 1968; Collins
and Urness, 1981; Mayle et al., 1999). Pellet groups were
counted every year in March along 2 m wide transects. Because
the disappearance rate of summer and autumn pellets is relatively
rapid in Poland (Aulak and Babińska-Werka, 1990b), all data
refer to patterns of winter habitat use by red and roe deer. The
transects were drawn on a 1:50000 map, so that their spatial
distribution in the study area was more or less even. Each transect
started at a characteristic point (e.g. a crossroads), which could be
easily found in the field. The bearing of each transect was
measured on the map and then walked with a compass. In total
there were eight transects. Transects walked in 1997 differed
slightly from subsequent years, when transects were drawn such
that they could be reached by car (using forest and not the main
470 J. Borkowski, J. Ukalska / Forest Ecology and Management 255 (2008) 468–475

Table 1 be preferred, e.g. grasses, raspberry (Rubus ideus), bramble (R.

The length of transect units (km) in different habitat types in year 2000
Total Palatable Unpalatable
T 2.7 – –
PS 3.3 – –
MF 26.0 – –
De 2.2 – –
P0 2.1 0.8 0.9
P1 4.0 1.2 2.4
P2 9.0 3.0 3.1
P3 8.7 2.7 2.5
T: thicket; PS: pole size forest; MF: mature forest; De: deciduous forest; P0:
pine forest with no understorey; P1: pine forest with sparse understorey; P2:
pine forest with medium understorey; P3: pine forest with dense understorey;
Palatable: patches dominated by palatable plants; Unpalatable: patches domi-
nated by unpalatable plants; Pal/Unpal: patches with both palatable and
unpalatable plants.
roads). Total transect length was about 32 km. Each year transect
length could differ slightly, for reasons such as recent forest work
in a given sub-compartment (e.g. tree cutting and young
plantation fencing) or high groundwater level.
Pellet groups were counted separately for each part of the
transect crossing an area of similar habitat characteristics (stand
age class, tree species, understorey and ground vegetation
condition). Each such part of the transect was considered a
sampling unit, hereafter called a transect section. The length of
each section was measured from the map or estimated by
pacing. The total number of sections in different study years
ranged from 108 to 184. Table 1 shows the length of sections in
different habitats. Average section length was 220 m, while the
average number of sections per transect was 18.4.
Beside the numbers of red and roe deer pellets alongside
transects, section data were collected on habitat characteristics
with regards to dominant tree species (pine, deciduous), forest
age class (thickets: 16–25; pole stage forests: 26–50; mature
forests: >50-year-old stands) and understorey or ground
vegetation conditions. Understorey and ground vegetation
conditions were recorded in mature pine stands only, since both
were scarce or absent in the other habitats. Understorey (woody
and shrubby vegetation up to 4 m in height) tree/shrub species
data (spruce, deciduous) was augmented by density informa-
tion. The understorey in each transect section was assigned to
one of four classes: no understorey (P0), sparse – composed of
single trees or shrubs (P1), medium-density – visibility distance
longer than 100 m (P2) and dense – visibility distance below
100 m (P3). Thus, the term ‘understorey density’ in this study
refers to visibility rather than to shrub/tree stem density. In
addition, understorey was classified as deciduous or spruce,
depending on composition. Although the classification of
understorey density was in this sense somewhat arbitrary, the
fact that a single observer collected most data (with only limited
help from another person) assured consistency.
Data on ground vegetation (herbaceous vegetation and
dwarf shrubs such as berries) was also collected within each
transect section. Based on food preferences of red and roe deer
(Dzie˛ciołowski, 1969; Ge˛bczyńska, 1980), the ground vegeta-
tion was classified as palatable (dominated by plants known to
caesius) and bilberry (Vaccinium myrtillus) or unpalatable
Pal/Unpal (dominated by plants which are usually avoided or seldom
– consumed, e.g. small-reed (Calamagrostis sp.) and sedges
– (Carex sp.)). For clarity, analysis was confined to those sections
– in which the ground vegetation could be clearly classified as
– dominated by palatable or unpalatable plants (more than 50%
0.4
of ground cover as estimated visually).
0.4
2.9
3.5 4. Statistical methods
Generalized linear mixed models GLMM (Breslow and
Clayton, 1993; Littel et al., 1996) were applied to the data for
each species separately. Two mixed models were used. The first
basic model (Model 1) was used to examine whether habitat
type, year and their interaction, influence forest use in red and
roe deer (habitat is defined as transect sections with similar
stand age and species, understorey and ground vegetation). The
second model (Model 2) analysed whether deer utilization of
mature forest with various understorey densities depends on
ground vegetation and its interaction with the understorey.
Pellet group count data per section was a linear combination
of fixed and random effects described by the generalized linear
mixed Model 1:
hi jk ¼ logðli jk Þ ¼ logðsÞ þ m þ yi þ h j þ ðyhÞi j þ tk (1)
where hijk is the log link function log(lijk) with a Poisson error
term, lijk the conditional mean (Littel et al., 1996) for the ijkth
year–habitat–transect combination, m the intercept, yi the ith
year fixed effect, hj the jth habitat type fixed effect, (yh)ij the
year–habitat interaction and tk is the random effect of the kth
transect. The logarithm of the section area, log(s), is an offset
term to allow for the different lengths of the transect sections.
Model 2 exclusively used data from mature pine forest where
ground vegetation was observed. The form of this model was
hi jkl ¼ logðli jkl Þ ¼ logðsÞ þ m þ yi þ u j þ gl þ ðugÞ jl þ tk
(2)
where lijkl is the conditional mean for the ijklth year–under-
storey–ground vegetation–transect combination, uj the jth
understorey fixed effect, gl the lth ground vegetation fixed
effect and (ug)jl is the understorey–ground vegetation interac-
tion and the other effects are the same as in Model 1.
Because empirical count data are typically over-dispersed,
(i.e., the variance is greater than the mean; Littel et al., 1996),
the significance of fixed effects of both models was based on the
F-statistics adjusted for over-dispersion. Degrees of freedom
were estimated using the Satterthwaite approximation, which
could give non-integer values. Differences between factorial
combinations were tested using Tukey–Kramer adjustment for
unbalanced data (Kramer, 1956).
In order to check if cover was equally important in habitat
used by the two deer species, index of selectivity was
calculated. Index of selectivity was defined as the ratio of
mean pellet group count values per area unit in examined
habitats to the general mean pellet group count value per area
 J. Borkowski, J. Ukalska / Forest Ecology and Management 255 (2008) 468–475 471

unit. The assumption was that the species with more

pronounced cover requirements will ‘‘prefer more’’ covered
habitats and ‘‘avoid more’’ open ones, while the species with
less pronounced cover requirements will not show these
preferences. As a result, the range of the index will be larger for
the former than for the latter.
Correlation between pellet group count data of the two deer
species was checked using the Spearman rank correlation. For
each species, transect residuals from Model 1 were compared
by Pearson’s correlation to see if there was any unexplained
variation correlated in the same way for red and roe deer. Both
correlations were done on the scale of the study area (sections
from all the transects together) and on the habitat scale
(separately for each habitat type).
Both GLMM models were fitted using the GLIMMIX
macro-implement in the SAS System for Windows 9.1.3 (SAS
Institute, 2002). Pearson’s correlation was assessed using the
CORR procedure of the SAS System.
5. Results
In total, 2890 red deer and 3101 roe deer pellet groups were
recorded. Red deer pellet group density differed significantly
among the habitats (F = 16.14; d.f. = 9, 540.4; p < 0.001) and
years of the study (F = 2.74; d.f. = 3, 543.7; p = 0.04), but was
not influenced by the interaction habitat  year (F = 0.71;
d.f. = 27, 538.6; p > 0.05). Similarly, habitat type was an
important factor influencing roe deer pellet group density
(F = 9.27; d.f. = 9, 545.3; p < 0.001). Roe deer pellet group
density was similar among the years (F = 1.46; d.f. = 3, 541.4;
p > 0.05); interaction habitat  year (F = 1.31; d.f. = 27, 542;
p > 0.05).
It was possible to rank the habitats based on mean pellet
group density (Fig. 1):
red deer : D  PS  P1d  P0  P1s  T  P2d  P2s
 P3d  P3s
red deer : D  P1d  P0  PS  T  P1s  P2s  P2d
 P3d  P3s
Fig. 1. Pellet group density means for red (closed bars) and roe (open bars) deer
per 100 m2 in different habitats. T: thicket; PS: pole stage forest; D: deciduous
forest; P0: pine forest with no understorey; P1: pine forest with sparse under-
storey; P2: pine forest with medium understorey; P3: pine forest with dense
understorey; s: spruce understorey; d: deciduous understorey. Error bars show 1
S.E.
For both species, the rankings were nearly identical and the
score of given habitat type depended very much on its cover
conditions. More meaningful however, are the comparisons of
specific pairs of habitats (Table 2). For both deer species, pellet
group densities were higher in mature pine stands with dense
deciduous and spruce understorey (P3d and P3s), and to some
extent in mature pine stands with medium density spruce
understorey (P2s; p < 0.05 or less), than in most other habitat
types (thickets, pole-sized and deciduous stands). In general,
there were no differences in either red or roe deer pellet group
densities among mature pine stands with sparse or no
understorey. Pellet group densities of both species were not
dependent on the kind of understorey (deciduous vs. spruce) in
any of understorey density classes ( p > 0.05 in all cases,
Table 2).
In mature pine stands, in all understorey density classes
except for the sparse understorey, there were significantly more
red deer pellet groups in transect sections with the ground
vegetation dominated by palatable plants then in those
dominated by unpalatable plants ( p < 0.05 or less, Fig. 2).
In the case of roe deer, more pellet groups were recorded in
sections with palatable plants than in unpalatable ones only in
P0, P2s and P3d ( p < 0.05 or less, Fig. 2).

Table 2
Significance of comparisons of red (bold font) and roe deer (underlined font) pellet group densities recorded in different habitats
T PS D P0 P1d P1s P2d P2s P3d P3s
T – ns ns ns ns ns ns ns * ***
PS ns – ns ns ns ns ns *** *** ***
D ns ns – ns ns ns ns/* * *** ***
P0 ns ns ns – ns ns ns * *** ***
P1d ns ns ns ns – ns ns ns ns ns
P1s ns ns * ns ns – ns *** *** ***
P2d ns ns * ns ns ns – ns ns ns
P2s ns * ** ns ns ns ns – ns ***
P3d ** *** *** ** ns ns/* ns ns – ns
P3s *** *** *** *** ns * ns * ns –
T: thicket; PS: pole stage forest; De: deciduous forest; P0: pine forest with no understorey; P1: pine forest with sparse understorey; P2: pine forest with medium
understorey; P3: pine forest with dense understorey; s: spruce understorey; d: deciduous understorey; ns: not significant; ns/*: 0.05 < p < 0.1, *p < 0.05, **p < 0.01,
***p<0.001.
472 J. Borkowski, J. Ukalska / Forest Ecology and Management 255 (2008) 468–475

Table 4
Correlation between red and roe deer pellet group densities in different habitats
Habitat type RS P
T 0.45 *
PS 0.46 *
De 0.51 *
P0 0.27 ns
P1d 0.22 ns
P1s 0.51 *
P2d 0.41 ns
P2s 0.29 ns
P3d 0.71 *
P3s 0.52 *
All habitats 0.57 *
T: thicket; PS: pole stage forest; MF: mature forest; De: deciduous forest; P0:
pine forest with no understorey; P1: pine forest with sparse understorey; P2:
pine forest with medium understorey; P3: pine forest with dense understorey; s:
spruce understorey; d: deciduous understorey; RS: Spearman correlation coeffi-
cient; P: probability; ns: not significant; *p < 0.05.

positive but insignificant ( p > 0.05). Similarly, there was a

Fig. 2. Pellet group density means and standard errors for red and roe deer per
100 m2 in transect sections dominated by palatale (closed bars) and unpalatable
plants (open bars) in mature pine forest: P0: pine forest with no understorey; P1:
pine forest with sparse understorey; P2: pine forest with medium understorey;
P3: pine forest with dense understorey; s: spruce understorey; d: deciduous
understorey; ns: not significant. *p < 0.05, **p < 0.01 and ***p<0.001.
positive correlation between the two sets of GLMM residuals
on the scale of the study area (r = 0.44 and p < 0.001, Table 5).
Significant ( p < 0.05 or less) positive correlations between the
residuals were recorded in 7 out of 10 habitats (Table 5).
6. Discussion

6.1. Pellet count credibility

The index of selectivity (Table 3) for red was more than one
and half times larger than for roe deer (1.88 and 1.17,
respectively). Similarly, the coefficient of variation of the index
for red deer was nearly twice as large as the one recorded for roe
deer (39% and 65%, respectively, Table 3).
On the scale of the study area, red and roe deer pellet groups
were similarly distributed (RS = 0.57 and p < 0.05, Table 4).
Pellet groups of both species were positively correlated in 6 out
of 10 habitats (Table 4), while in 4 remaining ones RS were
Dissimilarities in structural characteristics among habitats in
this study may have influenced deer pellet group density in
different habitats. Pellet decomposition rates may depend on
habitat conditions (Aulak and Babińska-Werka, 1990b),
particularly openness (Welch et al., 1990; Lehmkuhl et al.,
1994; Massei et al., 1998). However, in each of the cited
studies, habitat-related differences in decomposition rates were
minor in winter. Thus, it is unlikely that this factor exerted an

Table 5
Table 3 Correlation between the two sets of GLMM residuals (Model 1) for red and roe
Index of selectivity (see Section 3 for details) in red and roe deer habitat use deer in different habitats
Habitat Red deer Roe deer Habitat type RS P
T 0.94 0.83 T 0.47 ***
PS 0.49 0.75 PS 0.52 ***
D 0.25 0.41 D 0.41 *
P0 0.54 0.72 P0 0.47 **
P1d 0.50 0.68 P1d 0.02 ns
P1s 0.61 1.02 P1s 0.18 ns
P2d 1.29 1.29 P2d 0.35 ns
P2s 1.32 1.15 P2s 0.35 ***
P3d 1.93 1.58 P3d 0.65 ***
P3s 2.13 1.58 P3s 0.47 ***
Range 1.88 1.17 All habitats 0.44 ***
CV (%) 65 39
T: thicket; PS: pole stage forest; De: deciduous forest; P0: pine forest with no
T: thicket; PS: pole stage forest; De: deciduous forest; P0: pine forest with no understorey; P1: pine forest with sparse understorey; P2: pine forest with
understorey; P1: pine forest with sparse understorey; P2: pine forest with medium understorey; P3: pine forest with dense understorey; s: spruce under-
medium understorey; P3: pine forest with dense understorey; s: spruce under- storey; d: deciduous understorey; R: Pearson correlation coefficient; P: prob-
storey; d: deciduous understorey; CV: coefficient of variation. ability; ns: not significant; *p < 0.05, **p < 0.01 and ***p<0.001.
 J. Borkowski, J. Ukalska / Forest Ecology and Management 255 (2008) 468–475
important effect. Similarly, ground vegetation is scarce at the
end of winter and should not have much influence on the ability
to detect pellet groups. Ground vegetation therefore, was
assumed not to influence the results of this study.
6.2. Mature pine stand use
Our results suggest that mature pine forests (especially some
patches within them), were much more attractive to deer, than
has been reported for mature stands with spruce (Staines and
Welch, 1984; Catt and Staines, 1987; Latham et al., 1996). The
reason for this is probably poorer food conditions under the
spruce canopy, due to the presence of much less favourable
light conditions than in mature pine stands (Robakowski et al.,
2004). Mature spruce forests with ground vegetation have been
shown to sustain greater use on the part of red, and especially
roe, deer, than those lacking ground vegetation (Welch et al.,
1990). However, in this study, reference to the ground
vegetation alone could not account for the use deer made of
mature stands. For instance, use of deciduous stands offering
little food was similar to that of mature pine stands with little or
no understorey, that had much more abundant food resources.
6.3. Influence of security cover and ground vegetation
As predicted, cover seemed to be a very important
determinant of winter habitat use by the red and roe deer in
Rudy Raciborskie. General habitat rankings for both species
very much reflected a gradient of hiding cover conditions
offered by various habitats. Deciduous mature stands received
very little use, pole stage stands and mature pine stands without
or with sparse understorey received moderate use, and mature
pine stands with dense understorey were used most intensely.
When the number of visitors to an area is high, deer seek dense
cover (Jeppesen, 1987) even in populations generally tolerant
of human presence (Borkowski, 2001). As already mentioned,
this study area is commonly used, not only by people for
recreation, but also for hunting. Hunting promotes intensive
cover use by deer (Lovaas, 1958; Skolvin, 1982; Kufeld et al.,
1988). However, cover alone was not able to explain habitat use
pattern by deer in this study. For instance, there was no
significant difference in habitat use intensity between thickets
offering favourable cover conditions, pole stage stands with
neither food nor cover, or mature pine stands without
understorey which had abundant food but no cover.
Thus, the combination of food and cover seemed to
determine habitat attractiveness. Mature pine stands with a
dense understorey were the habitat type most preferred by both
deer species. No other habitat within the study area could have
offered both abundant food and good cover conditions. It has
been documented that simultaneous access to food and cover
can influence home range size. That is, the more fine-grained a
habitat mosaic, the smaller the home range that can encompass
all required resources (Moe and Wegge, 1994; Borkowski and
Furubayashi, 1998). Henry (1981) concluded that forest rides
were the habitat type most used by roe deer because of high
food availability and proximity to cover.
473
Although a dense understorey offering cover could also
attract deer through its abundant browse, this probably was not
the main reason for the use of such patches. Spruce twigs are
usually avoided by deer in Poland (Dzie˛ciołowski, 1969;
Borowski and Kossak, 1975), but the sites with a deciduous
understorey of higher quality browse were used as often as
those with spruce-dominated understories. Although Borowski
and Kossak (1975) found spruce bark to be important in winter
diets, in our study area we observed no spruce de-barking.
6.4. Comparison of habitat use between red and roe deer
6.4.1. Importance of ground vegetation
Ground vegetation was an important factor modifying deer
habitat use in Rudy Raciborskie. In case of red deer, this trend
was recorded in nearly every understorey density class and use
intensity was positively related to understorey density. Thus,
the cover seems to be a necessary primary condition for
preferred winter habitat of the studied deer species. Only when
cover is sufficient will food availability influence deer
distribution within this habitat. Although we do not have data
on animal activity in the studied habitats, these results suggest
that cover is important not only for resting but also for foraging
animals.
The use of mature pine stands by roe deer seemed to be less
influenced by ground vegetation than was found for red deer.
However, this is probably a result of the coarse classification of
patches with palatable or unpalatable plants. Roe deer is a
concentrate selector (Hofmann, 1985). The classification of the
patches in the present study probably better reflected the
conditions for an intermediate or roughage eater such as red
deer (Hofmann, 1985). Contrary to roe deer, red deer depends
more on food biomass than on its quality. In addition, unlike red
deer, roe deer winter fat reserves are relatively small and the
principal source of winter energy is food supply (Holand et al.,
1998). Therefore, winter habitat use by roe deer should have
been more influenced by food resources than was the case for
red deer.
6.4.2. Importance of security cover
In spite of similar habitat use patterns, roe deer (the smaller
species) seemed to find it easier to satisfy its cover requirements
than did the larger red deer. The idea of the more pronounced
cover requirements for red deer than for roe deer was supported
by wider ranges of the selectivity index and higher coefficient
of variation for red than for roe deer. A similar conclusion was
drawn by Lyon and Jensen (1980), who found that habitat use
among the smaller mule deer was less influenced by security
cover than was that of the larger elk. An especially clear
relationship between species size and the need for cover was
found in the relatively open post-burn area of young pine stands
at Rudy Raciborskie (Borkowski, 2004).
6.4.3. Interspecific relations
Latham et al. (1996) found that the densities of both deer
species in different forests of Scotland were negatively
correlated. In a subsequent study, Latham et al. (1997) pointed
474 J. Borkowski, J. Ukalska / Forest Ecology and Management 255 (2008) 468–475
out that there were several climatic factors (such as
temperature, snow depth, windspeed and rainfall), shaping
the negative relation between red and roe deer densities.
Moreover, within individual forests studied by Latham et al.
(1997), red and roe deer used their habitats in different ways. In
this study, densities of red and roe deer pellet groups were
positively correlated on the scale of the study area and within
many among available habitats. In winter especially, this should
not come as a surprise, since major differences in the feeding
habits of the two species observed in summer are likely to
disappear in winter (Staines et al., 1985; Latham et al., 1999).
Therefore, winter habitat use by both species may differ less
than in the other seasons. Similar habitat use patterns between
both species were also found in other studies (Staines and
Welch, 1984; Welch et al., 1990). Latham et al. (1997)
suggested that there may be an interspecific influence of red on
roe deer. In the present study, red and roe deer pellet group
densities were not negatively correlated, either on the study
area or the habitat scale. Nevertheless, there may be negative
effect of red on roe deer performance (body mass, survival and
reproduction), even with similar habitat preferences. As a
result, temporal exclusion or declining body mass of roe deer
could have occurred. During last decade however, no decline in
roe deer population number (Nasiadka, unpubl. data) or body
mass (Borkowski, in prep.) has been observed. In addition,
transect section residuals from the fitted models were positively
correlated for red and roe deer. This suggests that untested
factors were affecting both species in the same way. In
combination with similar habitat preferences, these results
suggest that interspecific competition was not an important
factor in this study. Nevertheless, the relations between the two
main and largely sympatric European deer species should
receive more attention from the researchers.
7. Conclusions
In hunted areas, security cover is an important factor
determining habitat use by deer, even though large predators are
absent. This study suggests that food also plays an important
role, but only where cover requirements are already met. Where
managers are interested in high densities of deer, it is important
to make sure that security cover is available. This is especially
true in habitats with abundant food and probably in areas where
people hunt in groups using beaters. Under such circumstances,
dense cover not only impedes shooting, but also allows some
deer to remain undriven. This issue should be considered,
especially in regions inhabited by larger deer species, since
cover requirements in smaller deer seem to be less pronounced.
Mature pine forests can serve as high-quality winter habitat
for deer, but only when they offer abundant understories. The
precise species composition of these understories is less
important in central Europe. The introduction of such
understories by foresters would thus seem important, not only
for general biocenotic reasons, but also on account of the ‘‘side
effect’’ of improved deer habitat. Such work can be seen as an
integrated forest and game management practice worthy of
promotion. However, in areas where large deer species cause
remarkable damage to forests, young plantations near forest
patches offering security cover may be under higher pressure
than those surrounded by poor cover stands.
Acknowledgements
We are grateful to the Headquarters of the Rudy Raciborskie
Forest District for assistance and the friendly atmosphere.
Thanks also to Dr. Pawel Nasiadka for his help and suggestions.
Prof. Rory Putman kindly commented on an earlier draft of the
manuscript.
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