Cent. Eur. J. Biol. • 8(1) • 2013 • 8-17
DOI: 10.2478/s11535-012-0108-2
Central European Journal of Biology
Autumn-winter diet overlap of fallow, red, and roe
deer in forest ecosystems, Southern Poland
Research Article
Artur Obidziński1,*, Piotr Kiełtyk2, Jakub Borkowski3,4, Leszek Bolibok5, Karolina Remuszko3,6
Department of Forest Botany, Warsaw University of Life Sciences,
02-776 Warsaw, Poland
1
Faculty of Biology and Environmental Sciences,
Cardinal Stefan Wyszyński University,
01-938 Warsaw, Poland
2
Department of Forest Ecology, Forest Research Institute,
05-090 Sękocin Las, Poland
3
Department of Forestry and Forest Ecology,
University of Warmia and Mazury,
10-727 Olsztyn, Poland
4
Department of Silviculture, Warsaw University of Life Sciences,
02-776 Warsaw, Poland
5
Forest Directorate Wielbark,
12-160 Wielbark, Poland
6
Received 30 May 2012; Accepted 21 September 2012
Abstract: The wild population of fallow deer in Central Europe has grown considerably over the last decade. However, information on feeding
habits of this alien species in relation to the indigenous red deer or roe deer, in areas of their co-occurrence, is scarce. A prevailing
view maintains that their food-niches are distinct, although direct comparative studies have not been carried out. Therefore, the aim
of the research was to compare the diets of fallow, red, and roe deer feeding in the same habitat. Research was based on the rumen
contents of 242 animals hunted in the autumn-winter season in the forests of Southern Poland. The analyses demonstrated that
fallow deer are moderate grazers in such conditions and eat more graminoids in comparison to red or roe deer (36.4% vs. 16.1% or
5.5%, respectively). On the other hand, it feeds on less browse (17.2% vs. 41.4%) or dwarf shrubs (8.4% vs. 19.0%) than red deer,
and on less bramble (10.9% vs. 34.6%) or forbs (4.0% vs. 7.6%) in comparison to roe deer (P=0.05). Although the diets of the
three deer species differ in terms of the proportion of each food type in their diet, overlapping of their food-niches is high (52.6%).
Keywords: Ungulates • Feeding preferences • Dama dama • Cervus elaphus • Capreolus capreolus • Central Europe
© Versita Sp. z o.o.
1. Introduction
Ungulates transform inhabited ecosystems to a great
degree. They influence species composition and
structure of plant communities [1-3]. They also have an
indirect impact on populations of invertebrates, birds,
and even small mammals [e.g. 4-6]. Therefore, the
significant growth of the number of ungulates recorded in
Europe [7,8] requires adequate examination, monitoring,
8
and regulation due to their role in the environment
[e.g. 9-11].
Roe deer, Capreolus capreolus, and red deer, Cervus
elaphus, are the most common cervids in Europe. Their
ecology is researched and presented in various studies
[e.g. 12,13]. Fallow deer, Dama dama, have been
historically less common and less well-studied. This
species was introduced in southern Europe from Asia
Minor in antiquity and is presently spreading intensely
* E-mail: [email protected]
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A. Obidziński et al.
In response to the aforementioned factors, we
focused our research on the study of the autumn-winter
diet of fallow deer, in comparison with the diet of roe and
red deer, in the forest environment of Central Europe.
The following hypotheses were formulated based on the
premises listed above: 1) graminoids constitute the main
component of fallow deer autumn-winter diet in central
Europe; 2) percentage share of particular food types in
the diet of fallow deer differs from that in the diets of roe
or red deer, so the overlap is small; 3) the diet of fallow
deer is more opportunistic, so it is more varied than that
of roe or red deer.
2. Experimental Procedures
2.1 Study area
The research was carried out in the Raciborskie Forest
over an area of ~40 000 ha situated in southern Poland:
N50º08’00”-23’10” and E18º11’50”-33’40”, at an altitude
of 178–252 m above sea level (Figure 1). Arenosols
prevail in the southern and western portions of the area.
They represent coniferous forest habitats with Querco
roboris–Pinetum potential vegetation. Cambisols prevail
in the north and east, and they represent broadleaved
forest habitats with Galio–Carpinetum and Melico–
Fagetum potential vegetation (Matuszkiewicz, J. M.
2008. “Potential natural vegetation of Poland – numeric
map”, IGiPZ PAN, Warszawa, http://www.igipz.pan.pl/
KĘDZIERZYNKOŹLE
ra
Od
er
Riv
across Central Europe [14]. For example, the number of
fallow deer has risen in Poland nearly fourfold in the years
2000–2010 – from approximately 6500 to 23300, in spite
of a simultaneous increase of the hunting harvest from
1700 to 5100 specimens yearly (Central Statistical Office.
Forestry 2010, http://www.stat.gov.pl/gus/5840_1540_
PLK_ HTML.htm, Last access 15.04.2012).
The growth of the fallow deer population in Europe
and its subsequent impact on the environment,
especially on forests, has resulted in keen scientific
interest. Results of a number of studies suggest little
impact of fallow deer on tree stands [e.g. 15-18]. while
others demonstrate significant economic damage and
unfavourable environmental changes [e.g. 19-23],
including inhibition of the natural regeneration of stands
and changes in the direction of vegetation succession
[e.g. 24,25]. Fallow deer most frequently feed on
graminoids [15,17,26-32]. However, occasionally their
diet is dominated by tree browse [19,23,28,30], acorns
[15,16,33], dead leaves and lichens [34], or fruits [35].
As fallow deer are an alien species in Central Europe,
it is of further value to establish whether there may be food
competition between fallow deer and indigenous cervids,
and if so, to what extent. This is especially important
considering observations that indicate that the growth
of the fallow deer population may cause a decrease in
the roe deer population [36,37]. Comparative studies
of the diets of the above-mentioned cervids inhabiting
the same areas simultaneously are scarce. They
demonstrate that fallow deer eat less forbs or browse
and more graminoids than roe deer [23,27], as well as
more graminoids, forbs, or acorn and less browse than
red deer [28,32,38]. However, no comparative studies of
all three – fallow, roe, and red deer – living in the same
place at the same time have previously been published.
Existing information on the fallow deer’s diet and its
comparison with roe or red deer’s diets comes mostly
from areas with mild winters [e.g. 15,16,19,28,33].
Snowy winters usually cause an increase in feeding
on browse in all cervids [e.g. 28,39,40]. However, even
seasonal differences in nutritional content in plants can
cause alter the composition of the diet [41]. Long lasting
snow cover, typical for the study area in Central Europe,
can cause different feeding behaviour in fallow deer from
that described in the south and west of the continent.
It can be supposed that limited access to forest floor
vegetation, due to a thick layer of snow, should result in
an increase of browse in the fallow deer’s diet, and more
intense competition between the three cervids species
for access to food supplies. Results of our study of the
fallow deer’s diet can be of practical significance for
decision making on this species’ population size control
in this part of the continent.
N
RUDY
W
E
S
POLAND
0
5
10km
research area
multiaged managed forest
main rivers
even-aged thickets
Figure 1.
Location of the study area.
9
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Autumn-winter diet overlap of fallow, red, and roe deer
in forest ecosystems, Southern Poland
analysed. Material for analyses was collected during
the September–February hunting season from the years
2004/5 to 2010/11. 242 rumina of hunted animals were
used for the study. Collected material included 84 fallow
deer, 93 red deer, and 65 roe deer. 127 specimens
were killed in autumn and 115 in winter. 86 samples
were obtained from large thickets and 157 samples
were obtained from regular managed forest (Table 1).
Samples were collected across the entire area of study
with a close to regular distribution. The distribution
of the sexes in each species sample was 37 males,
37 females, and 10 calves in fallow deer; 36 males,
38 females, and 19 calves in red deer; and 11 males, 48
females, and 6 fawns in roe deer. The lower number of
male specimens among the roe deer sample (caused by
hunting laws in Poland) does not need to be considered,
as differences between the diets of opposite sexes of
roe deer are generally insignificant [e.g. 39,45,46]. The
ages of hunted specimens were determined by dentine
layers, but was not always available as hunters did not
consistently described samples. Age was recorded for
54 out of 84 fallow deer, 42 out of 93 red deer, and 34
out of 65 roe deer specimens. Mean age of harvested
fallow deer was 4.14 years (±SD=2.00), while it was
2.29 years (±SD=2.36) for red deer, and 3.21 years
(±SD=1.41) for roe deer.
About 500 cm3 of the rumina’s contents was
collected directly after the specimens were killed, and
preserved in 5% formaline solution. Every sample
was labeled in the field. The date and location of
collection, as well as the species, gender and age
of each given specimen, was marked on the label.
Samples were then washed in a sieve of 1 mm mesh
in the lab. Subsequently, three samples of 5 cm3 each
were extracted from the fraction left in the sieve from
each rumen. They were then analysed using the ”pointframe” method [47], which involved mixing them with
water and smearing the samples on a tray with a grid of
lines forming 100 intersection points along the bottom,
allowing for random selection of 100 particles of food.
The procedure was repeated three times to obtain a
sample of 300 items, which is considered adequately
representative [48]. Collected particles were identified
geoekoklimat/roslinnosc/prn_mapa/home_pl.htm. Last
access 15.04.2012). The mean temperature is -1.2ºC
in January, and 18.1ºC in July. The mean annual
precipitation level is approximately 630 mm and the
vegetation period lasts about 224 days [42]. About
9000 ha of the Raciborskie Forest stands were burnt
in the year 1992. Over 6000 ha were replanted, mostly
with Scots pine. The remaining area regenerated on its
own, mainly with silver birch [43]. In the period of study,
regeneration reached the stage of thickets 11–18 years
in age.
Scots pine (Pinus sylvestris) dominates the forest as
a whole, occupying 68% of its area. Silver birch (Betula
pendula) occupies 13%, and pedunculate oak (Quercus
robur) occupies 9%. Norway spruce (Picea abies),
European beech (Fagus sylvatica), oak (Quercus robur),
and black cherry (Prunus serotina) prevail in the shrub
layer of the unburnt parts of the forest. Common bracken
(Pteridium aquilinum), bilberry (Vaccinium myrtillus),
reed grass (Calamagrostis epigeios), wavy hair grass
(Deschampsia flexuosa), quaking grass sedge (Carex
briziodes), and bramble (Rubus fruticosus) dominate
the forest floor. Pine with admixture of birch, European
larch (Larix decidua), and common aspen (Populus
tremula) are the most common types of vegetation in the
thickets overgrowing burnt areas. Reed grass, purple
moor grass (Molinia caerulea), wavy hair grass, and
common heather (Calluna vulgaris) [44] dominate the
forest floor of the thickets. Crops and canola, as well as
some corn and root crops, available from neighbouring
fields (personal observation). 150 tons of fodder beet,
approximately 100 tons of maize silage, about 100 tons
of crop grain (corn, oats, barley, and wheat), and roughly
30 tons of hay were distributed as winter feed (based
on oral information from local hunting clubs in 2010). In
2009, the populations of roe deer, red deer, and fallow
deer in the surveyed area were approximately 10.5,
14.8, and 9.6 individuals/km2, respectively (J. Borkowski
et al., unpublished data).
2.2 Sample collection and data analysis
Food obtained from the rumina of animals harvested
by hunters according to annual hunting plans was
Species
Red deer
Fallow deer
Roe deer
Season
autumn
winter
autumn
winter
autumn
winter
Forest
23
30
24
25
26
28
Thickets
18
22
26
9
10
1
Sum
41
52
50
34
36
29
Table 1.
Number of samples of categories of animals analyzed.
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A. Obidziński et al.
of food; 5) significance of each type of food in the diet;
and 6) overlapping of the species’ diets.
The niche breadth of food categories consumed
was assessed using Levins’ [51] measure: B=1/Σpi2,
where B is the niche breadth and pi is the proportion of
food category i in the diet. This index was standardized
to a scale of 0−1 following formula of Hurlbert [52]:
Bsta=(B–1)/(Bmax–1), where Bmax is the total number
of food categories consumed by at least one herbivore
species.
Similarities of samples were established with the
use of principal components analysis (PCA) based on
the correlation matrix of the variables [53]. Samples
obtained from the specimens’ rumina defined according
to percentage share of particular types of food were
used as statistical units. The PCA diagram (Figure 2A)
represents classification of specimens for each
species. A separate diagram (Figure 2B) represents
the contribution of each type of food for the specimens’
ordination in reference to PC1 and PC2. Factor loadings
expressing factor-variable correlation are presented in
Table 2. The absolute magnitude of their values reflects
the strength of the relationship between a variable and
particular axis. The signs + or – in their values indicate
positive or negative correlation of a variable with a
given axis. Calculations were done with the use of the
Statistica 8.0 package (StatSoft Inc. 2008. Statistica ver.
8.0).
Percentage share of each type of food in the diets
of the studied species were compared by using the
Kruskal-Wallis test with an adequate test of multiple
comparisons [54]. This method was chosen due
to different numbers of animals in each group and
impossibility of transforming data to normal distribution.
Implementation of the above-mentioned tests was done
by the naked eye or enlarged with a 25x magnifying
binocular. Particles were then divided into the following
groups: browse and needles of coniferous trees (Pinus
sylvestris, Picea abies, Larix decidua), browse and
leaves of broadleaf trees (mainly Prunus serotina and
Quercus robur), dwarf shrubs (Vaccinium myrtillus,
V. vitis-idaea, Calluna vulgaris), bramble (Rubus
fruticosus s.l.), grasses (mainly Deschampsia flexuosa,
Poa sp., Agrostis sp.), sedges (mainly Carex brizoides,
Carex ericetorum), dicotyledones forbs, mushrooms,
acorns, crops (mainly rape leaves), additional feed
given (mainly fodder beet and crop grains), pollution,
and unidentified. Plants collected in the study area
were used as reference material. Identified particles
were dried at 60°C for 48 hours and weighed with an
accuracy of ±0.001 g. Therefore, comparison of diets of
analysed species was based on dry biomass of food. As
food in each rumen was ruminated to a different degree,
the percentage share of a particular type of food in
the rumina was calculated based on dry mass of food
and not on frequency of food particles. In the cases of
strongly ruminated samples, the easily-digested forbs
made up a lower percentage of the mass than other
food types [49]. To compensate for these altered values,
some authors [e.g. 50] suggest doubling the amount
of forbs in the results. Forbs analysed in our research
were not doubled because samples were of varied
levels of digestion. It can be therefore assumed that the
actual percentage of forbs could be slightly higher than
demonstrated.
Diets of analyzed species of animals were
compared in terms of: 1) number of types of food; 2)
breadth of food-niche (diet diversity); 3) similarities of
sample contents and principal food types that samples
differed by; 4) percentage of dry biomass of each type
0
-2
Axis 2
Axis 2; eig.=1.423 (15.8%)
2
1
graminoids
dwarf shrubs
forbs
0
coniferous
browse
feed
deciduous
browse
acorn and
mushrooms
-4
-6
red deer
roe deer
fallow deer
-2
Figure 2.
cultivated plants
Axis 1; eig.=1.613 (17.9%)
0
2
bramble
Axis 1
-1
-1
0
1
Diversity of components of the autumn-winter diet of red deer and roe deer in the autumn-winter period in forest ecosystems (S Poland)
acquired with the PCA method; a) ordination of samples of rumina content, b) ordination of type of food.
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Autumn-winter diet overlap of fallow, red, and roe deer
in forest ecosystems, Southern Poland
Food
Axis 1
Axis 2
forbs
0.3591
0.1718
graminoids
0.5129
0.5295
bramble
-0.0409
-0.7776
deciduous browse
0.3792
0.0129
coniferous browse
-0.7882
0.1280
dwarf shrubs
-0.5402
0.3096
acorn and mushrooms
0.3017
-0.1919
feed
0.2560
0.0088
cultivated plants
0.0770
-0.5993
Table 2.
Results of principal components analysis (PCA); factor
loadings expressing correlations of variables (diet
components) with PCA axes.
with procedures from the agricolae library written by
de Mendiburu (2009, Agricolae: Statistical Procedures
for Agricultural Research. R package version
1.0-7. Vienna. Austria: The R Foundation for Statistical
Computing) for R package (R Development Core Team.
2008. R: a language and environment for statistical
computing. R Foundation for Statistical Computing.
Vienna. Austria. URL http://www.R-project.org, on-line
access 15.01.2008). Percentage share of mass of
chosen category of food in a single rumen examined
was used as statistical unit in calculations. The level of
significance in statistical analyses was set at α=5%.
Index of importance of food type in the diet was set
after Bruno and Apollonio [19]: W=100Ni/N x 100Mi/M,
where Ni = number of rumina where food i was found,
N = number of all rumina analysed, Mi = mass of food i in
all rumina, and M = mass of all type of food in all rumina.
This index indicates, in a most complete manner, the
significance of particular types of food for each species
of animal. However, it unfortunately does not allow for
establishment of the significance of differences between
obtained values.
Dietary overlap was calculated using Schoener’s
index [55]:
O=100(1-0,5∑|Xi-Yi|) [%],
where Xi and Yi are the proportions of each food
category in the rumina of species X and Y.
3. Results
Food type analysis demonstrated the presence of 2–11
out of 13 types of food in any particular rumen. The mean
(±SD) number of types of food was highest in the rumina
of red deer (8.1±1.2), lower in fallow deer (7.7±1.2), and
lowest in roe deer (7.3±1.8). Only differences between
roe deer and red deer were significant (Kruskal-Wallis
test P=0.05).
The food-niche breadth of the examined animals
is characterised by high variability (0.01–0.38). Fallow
deer and red deer demonstrated similar mean foodniche breadth (0.16±0.08 and 0.17±0.07, respectively)
and at the same time were significantly (Kruskal-Wallis
test P=0.05) higher than that of roe deer (0.12±0.07).
Graminoids and forbs were the types of food
that distinguished the diet of the fallow deer from the
other two species’ diets. Bramble was the distinctive
type of food for roe deer, and needles, coniferous
browse, and dwarf shrubs were distinctive for red
deer (Figures 2A,B). More differences were found
between fallow deer and roe deer than between red
deer and either of the other two species. Each axis in
the principal components analysis explained a limited
amount of the entire variability. Eigenvalues of the nine
extracted axes amounted to 1.6131, 1.4230, 1.2155,
1.1012, 1.0147, 0.9216, 0.8748, 0.8331, and 0.0031,
which correspond to the following percentages of total
variability demonstrated by each axis: 17.92%, 15.81%,
13.51%, 12.24%, 11.27%, 10.24%, 9.72%, 9.26%, and
0.03%, respectively. Axis 1 was the most negatively
correlated with coniferous tree browse and needles
(r=–0.788) and dwarf shrubs (r=–0.540), whereas it was
positively correlated with graminoids (r=0.513) (Table 2).
Axis 2 was the most negatively correlated with bramble
(r=–0.778) and crops (r=–0.599), whereas it was
positively correlated with graminoids (r=0.529).
Frequency of type of food identified in the rumina
of examined species was high, reaching on average
more than 50% (mean 59.2%) with the exception of
acorns (27.3%), mushrooms (10.3%), fodder supplied
by hunters (18.5%), and crops (4.5%), as well as, in the
case of fallow and roe deer, browse (38.6%). Graminoids
(97.8%), needles (92.2%), and dwarf shrubs (91.4%)
were found to be most abundant in the rumina of red
deer. Graminoids (97.6%), needles (97.6%), and forbs
(91.7%) were found to be most prevalent in the rumina of
fallow deer. Bramble (87.7%), graminoids (86.2%), and
needles (81.5%) were found to dominated the rumina of
roe deer. Dwarf shrubs and browse were found in higher
frequency in rumina of red deer than in the other two
species, whereas, mushrooms and crops were of higher
frequency in the rumina of roe deer than in the other two
species (Table 3).
The percentage share of dry mass of each type
of food in the diets of analysed species differed
significantly. Needles, dwarf shrubs, and graminoids codominated in the diet of red deer (20.3%, 19.0%, and
16.1%, respectively). Graminoids (36.4%) composed
the highest percentage share in the diet of fallow deer,
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Frequency
Food
% share of dry mass ± SD
Importance
Red
deer
Fallow
deer
Roe
deer
Red deer
Fallow deer
Roe deer
Red
deer
Fallow
deer
Roe
deer
forbs
86.0
91.7
75.4
1.7±2.4 b
4.5±6.6 a
7.6±15.2 b
74.8
308.5
367,8
graminoids
97.8
97.6
86.2
16.1±16.9 b
36.4±24.1 a
5.5±12.1 c
689.7
2842.3
263,2
bramble
82.8
86.9
87.7
9.0±13.1 b
10.9±14.5 b
34.6±29.7 a
380.2
578.1
2329,6
deciduous leaves
89.2
77.4
80.0
9.113.6 a
6.4±12.0 a
7.9±12.6 a
398
415.8
453,6
coniferous needles
92.5
97.6
81.5
20.3±18.9 a
10.7±13.9 b
14.4±22.1 b
1145
1000.2
1200,7
dwarf shrubs
91.4
78.6
76.9
19.0±21.1 a
8.4±13.6 b
10.9±18.1 b
986.8
531.7
914,8
deciduous shoots
58.1
46.4
43.1
5.9±11.8 a
3.0±8.4 b
1.6±4.1 b
206.6
104.7
85,1
coniferous shoots
64.5
28.6
38.5
6.1±12.9 a
2.3±6.1 b
2.8±5.3 b
323.5
78.2
111,2
acorn
21.5
31.0
26.2
3.3±11.5 a
5.2±11.5 a
8.3±22.6 a
105.1
213.7
601,5
mushrooms
3.2
8.3
20.0
0.5±5.0 b
1.0±4.9 b
1.8±7.5 a
0.5
5.8
34,3
feed
22.6
31.0
3.1
6.6±15.3 a
7.4±16.1 a
0.9±5.1 b
111.8
395.1
2,5
cultivated plants
3.2
1.2
9.2
0.5±3.1 ab
0.4±3.4 b
1.7±6.4 a
1.4
0.2
12,4
other
96.8
94.0
96.9
2.0±2.7 b
3.5±3.6 a
1.9±2.2 b
103
232.4
142,3
Table 3.
Frequency, percentage share, and importance of food types in the autumn-winter diet of fallow deer, roe deer, and red deer in forest
ecosystems (S Poland). Red deer N=93, fallow deer N=84, roe deer N=65. The same symbol of the alphabet is used for marking means
that do not differ according to the level of significance set as P=0.05 for the same type of food.
followed by a significantly lower percentage share of
bramble (10.9%) and needles (10.7%). The diet of roe
deer was dominated by bramble (34.6%), while needles
(14.4%) and dwarf shrubs (10.9%) demonstrated
significantly lower percentage share. Red deer ate more
needles, dwarf shrubs, and browse in comparison with
the other two species. Fallow deer ate more graminoids,
and roe deer ate more bramble, forbs, mushrooms, and
crops (Kruskal-Wallis test P=0.05; Table 3).
The value distribution of importance indices of types
of food was to a great degree similar to the percentage
share of food mass (Table 3). Needles (1145), dwarf
shrubs (987), and graminoids (690) reached the highest
values in the diet of red deer. Graminoids (2842) played
the most important role in the diet of fallow deer, followed
by still high though lower percentage share of needles
(1000) and bramble (578). Bramble (2330) dominated
the diet of roe deer, followed by the still important though
smaller role played by needles (1201) and dwarf shrubs
(915). In spite of significant differences in percentage
share of particular types of food in the diets of fallow
deer, red deer, and roe deer, 52.6% of the diet was
common between all three species. Dietary overlap
reached 70.4% between red deer and fallow deer, 59.7%
between fallow deer and roe deer, 60.9% between roe
deer and red deer (Figure 3).
roe deer
red deer
8.3
21.3
32.0
52.6
7.1
17.8
22.5
fallow deer
Figure 3.
Overlap of the autumn-winter diets of red deer, fallow
deer, and roe deer in forest ecosystems (S Poland),
based on mean percentage share of food mass.
4. Discussion
The diet of the fallow deer analysed is highly varied.
Although graminoids constitute the largest part (over
1/3) of the diet, they were less abundant than in the
cases where fallow deer behaved as typical grasseating species [e.g. 15,27-29,35]. On the other hand,
percentage share of browse in their diet was too small
13
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Autumn-winter diet overlap of fallow, red, and roe deer
in forest ecosystems, Southern Poland
to consider fallow deer a browser species – as was
demonstrated in a few other places [19,23,30,56].
Food supplies present in the study area described
in the literature [44] encompassed eight categories:
deciduous branches, coniferous branches, deciduous
seedlings, coniferous seedlings, dwarf shrubs, bramble,
forbs, graminoids, and mosses. The percentage share
of the dry mass of food found in the rumina was
compared with that of their share in the environment
with the Ivlev’s electivity index [57]. We found that
fallow deer preferred dwarf shrubs and bramble while
they avoided tree leaves, forbs, and mosses. They
ate graminoids and woody browse proportionally to
these food types’ share in the environment. The diet
of red deer analyzed in this study was rather varied
and had a medium percentage share of plants rich
in fibre. In comparison with the food acquired by red
deer in coniferous forests in other locations [58], it
contained more needles, woody browse, and bramble,
less forbs, and similar amounts of dwarf shrubs and
tree leaves. With respect to the food supply available
in the study area [44], red deer preferred dwarf shrubs
and avoided forbs and mosses. Red deer acquired
other types of food according to their prevalence in
the environment. The diet of roe deer analyzed in
this study was dominated by plants of high nutritive
value. In comparison to food acquired by roe deer in
coniferous forests in other locations [59], it contained
significantly more bramble, less dwarf shrubs and
graminoids, and a similar share of forbs and coniferous
browse. With respect to the food supply available in the
study area [44], roe deer preferred bramble and dwarf
shrubs and they avoided graminoids, tree leaves, and
mosses. They fed on needles and forbs accordingly
to their prevalence in the environment. Roe deer
demonstrated the highest selectivity while fallow deer
demonstrated the lowest. At the same time, the diet of
the fallow deer was more similar to that of the red deer
in terms of contents and variety than to the diet of the
roe deer. The impact of additional feeding and crops
available on neighboring fields on the composition of
the diet of all three species may be considered to be
small. The percentage share of cultivated plants in the
rumina of all three species did not exceed 2% and in
winter was close to zero. Although fallow deer are
often considered to be more eager to wander out of
the forest and onto fields than red deer, the diet does
not support this view. The percentage share of fodder
given by hunters did not exceed 7.5% of the diet of any
of the analysed species in the period of study, and in
winter it constituted a maximum of 10.5% of dry mass
of their food. It is worth mentioning that similar feeding
is practiced all over Poland.
Differences between the diets of the studied
species generally resemble results obtained by earlier
studies of diet in these species. Similar to the results
presented here, fallow deer were previously found to
feed on more graminoids and forbs and less browse
with respect to red deer [28,32,38]. On the other hand,
fallow deer demonstrated smaller consumption of dwarf
shrubs than the other two species. Also, similarly to
other authors, we found fallow deer to feed on more
graminoids and less forbs, browse, leaves, and needles
in comparison to roe deer [23,27]. However, contrary
to the findings of other authors, we recorded a lower
consumption of bramble, mushrooms, and crops, and
greater consumption of fodder provided by hunters, as
well as a lack of differences with respect to consumption
of browse. However, it should be emphasised that food
availability in a given environment has an important
impact on the results [58,59].
Differences in diets of fallow deer, red deer, and
roe deer seem to be a constant predisposition as they
are based on the animals’ different structure and the
functioning of the digestive tract. The digestive tract
of the roe deer best digests forbs, while fallow deer
- graminoids, and the red deer’s digestive system lies
somewhere in between these two [60,61]. It is further
supported by the fact that in spite of a close evolutionary
relationship between of these three species, the fallow
deer’s diet is sometimes more similar to the diet of
typical grazing species, such as bovines or even equids
[27].
The overlap of the autumn-winter diet of fallow
deer with that of indigenous cervids across the study
area was higher than in the west of the continent. It
was higher by 13.3% in the case of fallow deer and
roe deer than in England, as calculated from Putman’s
data [27]. In the case of fallow deer and red deer, it
was higher by 17.4% than in Spain, as calculated from
the data of Garcia-Gonzales and Curtas [28]. Differing
development of vegetation resulting from the diverse
climates of these areas must be taken into account in
the interpretation of these differences. The latter may
result from lower autumn-winter food diversity in the
environment with longer and harder winters (Poland). A
change in diet can be caused not only by lack of access
to certain food types in given seasons, but also to a
change in the nutritive properties of the same plants
over different seasons [41]. Both of these factors can
cause the overlap of food-niches between cervids to be
higher in winter than in other seasons [48,62]. Particular
climatic conditions may cause food competition between
fallow deer and indigenous cervids [62], especially in a
situation when populations are too high or winter lasts
longer than usual.
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A. Obidziński et al.
On the other hand, the overlap of the diet does not
necessarily result in interspecific competition. Given
that a preferred type of food is supplied in abundance,
two species can co-exploit it without detriment to one
another [63]. In order for exploitation competition to take
place, an overlap in habitat use and in diet must occur
simultaneously and the shared dietary resource must be
limited [e.g. 48,64,65].
Food competition between fallow deer and the other
two deer species has not been previously recorded in
Poland. However, in view of the rapid growth of the
fallow deer population, the question whether this status
quo will remain or not is valid. This concern is supported
by observations made in Italy of decreasing roe deer
populations in areas where fallow deer settled [36,37],
as well as observations suggesting the same from
England [66]. Food competition between red deer and
fallow deer is suggested by a number of studies [32,66].
Rapid growth of the fallow deer population gives cause
for concern in terms of imminent increase in damage in
young tree stands as well, after introduction of fallow
deer next to roe and red deer in Central European
forests. Although fallow deer generally avoid browse,
they feed on it during the winter.
The results of this study and those in the published
literature indicate that fallow deer feed on their
preferred food – graminoids – whenever possible. On
the other hand, they are a flexible species and adjust to
the food supply available in the environment. Overlap
of food niches between fallow deer and the indigenous
cervids is small in environments and seasons with
diverse food-plant supplies. Overlap of their food
niches increases in environments and seasons of low
diversity of these supplies. Competition can occur
between these species, and it may grow stronger in
Europe with increasing of geographic longitude and
latitude, especially with further growth of fallow deer
populations.
The results of this study provide reasons for setting
permissible densities of fallow deer populations in order
to prevent competition between this species and red or
roe deer in different ecosystems or climatic regions. It
is worth mentioning that setting the right numbers for
fallow deer populations should take into account the
body mass ratio of 1:2 of fallow deer to roe deer and 2:1
of fallow deer to red deer [67]. It also seems advisable
to leave a number of hunting areas without fallow deer
as control plots for monitoring the relations between this
species and the indigenous cervids co-inhabiting with
fallow deer in other locations.
Acknowledgements
We would like to thank hunters from the following
hunting clubs: Azoty, Daniel, Darz Bór, Łoś, Odyniec,
Ostoja, Ponowa, and Ryś, and particulary to Mr.
Ryszard Kot and Mr. Zygmunt Tomaszek for collecting
the rumen content samples. We are also grateful to
Prof. Jacek Goszczyński of Warsaw University of Life
Sciences for helpful suggestions to the first version of
this paper and to anonymous Referees for their valuable
and encouraging remarks. Research was financed
by the Faculty of Forestry WULS and by the General
Directorate of Polish State Forests with research grant
No. BLP-267. Translated by Bogna J. Obidzińska.
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