Forest Ecology and Management 407 (2018) 166–173

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Forest Ecology and Management

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Diversionary feeding can reduce red deer habitat selection pressure on T

vulnerable forest stands, but is not a panacea for red deer damage
⁎,1
Johanna Maria Arnolda,b, , Philipp Gerhardtc,1, Sam M.J.G. Steyaertd,e, Eduard Hochbichlerc,
Klaus Hackländera
a
Institute of Wildlife Biology and Game Management, Department of Integrative Biology and Biodiversity Research, BOKU – University of Natural Resources and Life
Sciences, Vienna, Gregor-Mendel-Straße 33, 1180 Vienna, Austria
b
Wildlife Research Unit of Baden-Württemberg, LAZBW, Lehmgrubenweg 5, 88326 Aulendorf, Germany2
c
Institute of Silviculture, Department of Forest and Soil Sciences, BOKU – University of Natural Resources and Life Sciences, Vienna, Peter-Jordan-Straße 82, 1190 Vienna,
Austria
d
Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life Sciences, 1433 Ås, Norway
e
Faculty of Technology, Natural Sciences and Maritime Sciences, University College of Southeast Norway, 3800 Bø i Telemark, Norway

A R T I C L E I N F O A B S T R A C T

Keywords: Diversionary feeding implies strategic food provisioning to wildlife to lure animals away from undesired areas,
Cervus elaphus and is a common forest management practice throughout Europe and North America. Within forestry, diver-
Supplementary feeding sionary feeding typically targets cervids, and aims to reduce browsing damage and bark-stripping in vulnerable
Diversionary feeding forest stands (young and mid-aged forest stands). Because these stands are most vulnerable during winter, di-
Browsing
versionary feeing is often restricted to that season. Despite being widely applied, clear evidence on the effec-
Bark-stripping
tiveness of diversionary feeding of cervids is lacking. The aim of this study was to evaluate the effectiveness of
Deer impact
diversionary winter feeding as a tool to lure away a large cervid, red deer (Cervus elaphus), from vulnerable forest
stands. First, we hypothesized (H1) that diversionary feeding is among the most important factors explaining
differences in red deer habitat selection between summer (i.e., no food supply at feeding stations) and winter
(i.e., food supply ad libitum at feeding stations). Second, we hypothesized (H2) that diversionary feeding releases
red deer habitat selection pressure in forest stands vulnerable for browsing or bark-stripping, and that this
release decays with distance to the nearest feeding station. We tested our hypotheses using red deer relocation
data (11 individuals & 3 years) and resource selection functions in an intensively managed Austrian forest. As
expected, variation in red deer habitat selection between summer and winter was best explained by a different
response to supplementary feeding stations between the seasons (H1). During winter, red deer strongly selected
for areas close to feeding stations, whereas feeding stations did not strongly affect their habitat selection during
summer. We found that diversionary feeding during winter released red deer habitat selection pressure on
vulnerable forest stands, but limited to distances of about 1.3 to 1.5 km from the feeding station, this accounts
for 39% of the study area (H2). Our results show that diversionary feeding can indeed be an effective tool to
mediate habitat selection behavior and to lure cervids away from vulnerable forest stands. However, habitat
selection is not necessarily a good proxy for damage, and whether or not diversionary feeding reduces forestry
damage in the long-term remains unclear, as it can positively feed-back into survival, reproduction, and even-
tually population growth and densities. We suggest that forest managers also consider silvicultural measures,
population control (i.e., hunting) or spatial planning as means to minimize red deer induced forestry damage.

1. Introduction sometimes unexpected ways (Putman and Staines, 2004; Robb et al.,
2008). Supplementary feeding can alter behavior, life history, and de-
Supplementary feeding wildlife is a common but controversial mography in wildlife (Boutin, 1990; Robb et al., 2008). Further, it can
management practice, because it can influence wildlife in many, affect species assemblages and distribution (Redpath et al., 2001),

⁎
Corresponding author at: Wildlife Research Unit of Baden-Württemberg, LAZBW, Lehmgrubenweg 5, 88326 Aulendorf, Germany.
E-mail address: johanna.arnold@lazbw.bwl.de (J.M. Arnold).
1
Equally contributing authors.
2
Present address.

http://dx.doi.org/10.1016/j.foreco.2017.10.050
Received 9 July 2017; Received in revised form 21 October 2017; Accepted 24 October 2017
Available online 07 November 2017
0378-1127/ © 2017 Elsevier B.V. All rights reserved.
J.M. Arnold et al.
social organization and intraspecific interactions (Craighead et al.,
1995), and it can enhance parasite and pathogen transmission (Bradley
and Altizer, 2007). Furthermore, supplementary feeding can mediate
human-wildlife conflict through food conditioning and human habi-
tuation (Kavčič et al., 2013; Steyaert et al., 2014).
Diversionary feeding implies strategic food provisioning to wildlife
to lure animals away from undesired places or habitats (Kubasiewicz
et al., 2016). It is common in forest management throughout Europe
and North America, and typically targets cervids (e.g., red deer Cervus
elaphus, moose Alces alces) to reduce browsing damage and bark-strip-
ping (Putman and Staines, 2004; van Beest et al., 2010). Its effective-
ness regarding reducing red deer damage on forests is not unambiguous
and appears to be context dependent (Kamsker, 1979; Ueckermann,
1986; Schmidt and Gossow, 1991; Náhlik et al., 2005). The underlying
reasons for the effectiveness of diversionary feeding of deer are not fully
understood as they are highly complex and depend on many factors,
such as habitat, winter severity, feeding site characteristics, as well as
type, quality and quantity of provided food and its arrangement at
feeding sites, as well as local history of its use (Pheiffer, 1983; Nopp-
Mayr et al., 2011; Jerina, 2012; Gerhardt et al., 2013; Ossi et al., 2017).
Red deer is an intermediate feeder with a tendency towards
roughage eater (Hofmann, 1989). From spring to autumn, red deer diet
consists predominantly of protein rich forage such as grasses and herbs
(Schröder, 1977; Trdan and Vidrih, 2008), and deer inflict little damage
to commercial forest stands (Gill, 1992). During winter, when grasses
and herbs lignify and become unavailable by snow and ice, twigs and
buds of trees and shrubs with high fiber content become important food
resources (Schröder, 1977). In alpine habitats, red deer prefer con-
iferous trees as foods during winter (e.g., silver fir Abies alba, Norway
spruce Picea abies) (Schröder, 1977), which are also commercially the
most interesting species. Hence, diversionary feeding is typically only
conducted during the winter months.
Red deer is a conflict-rich game species for forestry (Reimoser and
Reimoser, 2010). It can negatively affect forest productivity by
browsing or bark-stripping. For example, in Austria, herbivores (red
deer, roe deer Capreolus capreolus, and chamois Rupicapra rupicapra) can
cause damage by browsing and/or bark-stripping of up to about 218
million Euro per year in at least 10.000 km2 or 25% of the total forest
area per year (Reimoser, 2000; Reimoser and Putman, 2011). In addi-
tion, they cause minor damage to tree regeneration because of fraying
and trampling. By feeding on apical buds and shoots, tree growth can be
suppressed or be hampered, and red deer browsing can prolong or
pause forest cycles. Selective feeding affects species composition and
turnover (Lilleeng et al., 2016). By bark-stripping, fungi can enter the
wounds and destroy lignin, leading to economic losses and higher
susceptibility of trees to abiotic impacts such as snow damage or
windthrow. Unequivocally, by browsing and bark-stripping, red deer
has direct impacts on commercial trees and forest vegetation in general
(Gill, 1992; Reimoser et al., 1999). Red deer-induced forestry damage is
of growing concern for forest managers in Europe (Verheyden et al.,
2006; Jerina et al., 2008; Kiffner et al., 2008; Klopcic et al., 2010).
Diversionary feeding during winter is the most widely applied tool in
forestry and wildlife management to lure red deer away from vulner-
able sites (Putman and Staines, 2004). Red deer show an opportunistic
feeding strategy, selecting for easy available food resources in their
home ranges. Artificial food resources, such as supplementary winter
feeding, may alter red deer spatio-temporal behavior (Luccarini et al.,
2006; Jerina, 2012).
There is a lack of empirical studies examining the efficiency of di-
versionary feeding regarding browsing and bark-stripping. To our
knowledge, no study so far has tested the mechanisms of selection for
specific habitat types in relation to winter feeding schemes. Here, we
assess the effectiveness of diversionary feeding during winter to keep
red deer away from forest stands that are vulnerable for browsing and
bark-stripping. First, we compare red deer habitat selection between the
period of diversionary feeding (hereafter ‘winter’, when feeding stations
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Forest Ecology and Management 407 (2018) 166–173
are supplied with food ad libitum) and no diversionary feeding (here-
after ‘summer’, when feeding stations are not supplied with food). We
hypothesize (H1) that red deer response to feeding stations is the most
important factor that distinguishes red deer habitat selection between
winter and summer, and that red deer selects for areas close to sup-
plementary feeding stations during winter but not during summer.
Secondly, we zoom in on red deer habitat selection during winter, and
hypothesize (H2) that forest stands that are vulnerable for browsing or
bark-stripping become increasingly released from red deer habitat se-
lection pressure the closer to supplementary feeding stations. We test
our hypotheses based on red deer GPS relocation data and resource
selection functions (RSFs) in an intensively managed forest in Styria,
Austria.
2. Materials and methods
2.1. Study area
The study area is located in the Eastern Central Alps in the upper
part of the river Mur valley in the federal state of Styria, Austria. The
region is located between 46°59′12″ und 47°04′20″ latitude and
13°53′59″ and 14°00′27″ longitude, and covers a total area of 131 km2.
Average annual temperature in our study area is 5 to 6 °C and the
precipitation varies in average between 60 and 160 mm per month,
with minima during winter. Although precipitation is low during winter
months, mean temperatures below zero maintain a snow layer from
around December to March. Summer months are warm and wet.
Shielded valleys have a pronounced continental climate, whereas
overall the climate is milder inner alpine (Kilian et al., 1993). The study
area comprises a wide altitudinal gradient (850–2200 m.a.s.l.), and
habitat types range from submontane to high subalpine habitat. The
study area consists of a privately owned forest, managed for timber
production as well as for hunting, maintaining high densities of red
deer stags older than 10 years. Large carnivores, such as grey wolf
(Canis lupus) or Eurasian lynx (Lynx lynx) are observed only occasion-
ally (Erwin Lick, personal communication). Forest vegetation is char-
acterized by a transitional zone of silver fir (Abies alba) – Norway
spruce (Picea abies) forests with a predominance of the latter. In lower
altitudes, broadleaved species such as sycamore (Acer pseudoplatanus)
and European beech (Fagus silvatica) are admixed. The higher subalpine
consists mostly of European larch (Larix decidua) and stone pine (Pinus
cembra). Economically important is the harvest of Norway spruce and
European larch. Overall, the area is managed with a small clear-cut
regime and therefore has a patchwork-like structure, intersected with
forest roads. The study area suffered from heavy windthrows in 2002,
when about 7.3 km2 hectares of forest was completely wind-thrown.
These areas were afforested afterwards. These windthrows resulted in a
heterogeneous pattern of old forest stands and regenerating forests.
Red deer density varies between forest beats. According to si-
multaneous counts at the winter feeding stations of the forest en-
terprise, densities vary from 7 to 12 animals per 1 km2 (Lick, 2013), in
average 9 animals per 1 km2 (Arnold and Hackländer, 2014). Red deer
hunting season starts, depending on sex and age, on 15th May and lasts
until 15th January. Winter feeding occurs outside the growing seasons
and depends on weather conditions. Roughage (hay) is provided ad li-
bitum and daily complemented with succulent feed (mainly apple po-
mace) at seven active feeding stations (1 per 18.7 km2) in the study area
from October to May. Exact start and end of food provisioning de-
pended on onset of winter and start of growing season. Feeding stations
are located on clearings in thinned old growth forest stands, ranging
from 1680 to 1810 m.a.s.l. Damage induced by ungulates is at a critical
level: A comprehensive sampling plot procedure in 2009 and 2010
showed that about 33.5% of apical buds of spruce regeneration
(trees < 0.2 to < 1.80 m) being browsed annually, and the bark of
about 2.3% of all stems in spruce pole wood stands (aged 20–39 years)
being stripped from bark each year. Bark-stripping damage amounts to
J.M. Arnold et al.
21–30% of the total stem number per hectare of Norway spruce in
commercial forest stands and up to 45% in protection (e.g. for rock fall
or avalanches) forests. Browsing on spruce by herbivore ungulates af-
fects about 40% of the regeneration plants each year (Gerhardt and
Hochbichler, 2011).
2.2. GPS-relocation data
We captured and collared adult red deer with Global Positioning
System collars (GPS; Vectronic Aerospace GmbH, GPS PLUS collar).
Authorized veterinarians immobilize red deer from high seats at the
feeding stations with a remote drug delivery system. Immobilization
(with “Hellabrunner Mixture”, Wiesner, 1989) and radio-collaring
complied with current laws in Austria and Styria, and were approved by
the relevant authorities. Collaring took place during diversionary
feeding season in the years 2009 and 2010. Deer positions were re-
corded every three hours. GPS relocations were assorted according to
the two seasons, and according to the exact start and end of food pro-
vision per year and feeding station.
2.3. Landscape variables
Data regarding habitat type, elevation data (slope steepness, aspect,
and elevation), anthropogenic infrastructure, and feeding stations were
extracted from forest inventory maps from the landowner using the
software ArcGIS 10.4.1 (ESRI, 2016). We grouped habitat types
(10 ∗ 10 m) according to their vulnerability to browsing or bark-strip-
ping from a forestry perspective, and accounted for other habitat types
that likely affect red deer habitat selection. We considered young for-
ests (i.e., natural or artificial regeneration stands until canopy closure)
and windthrow areas (i.e., open areas > 0.3 ha wind-thrown in 2002,
no canopy closure) as two separate habitat types vulnerable for
browsing. We classified thickets (i.e., forest stand > 1.30 m until dia-
meter at breast height (dbh) of 10 cm) and pole wood stands (i.e.,
dbh > 10–20 cm) as bark-stripping vulnerable habitat. We merged
timber wood (i.e., dbh > 20–50 cm), old growth (i.e., dbh > 50 cm
and more than 150 years old), and krummholz stands (i.e., dwarf
mountain pine Pinus mugo above timberline) as ‘not vulnerable forest’.
We also extracted alpine meadows (i.e., alpine pasture with cattle
grazing), meadows (i.e., sown pastures and grasslands), and other non-
forest land (e.g. ski slopes, wasteland) as land-cover types within the
study area. We extracted altitude from a 20 ∗ 20 m digital elevation
model, and distance to the nearest creek and road (10 ∗ 10 m) form
digital topographic maps. We extracted distance to the nearest sup-
plementary feeding station based on forest inventory data from the
landowner.
2.4. Statistical analysis
We used resource selection functions (RSFs) to assess how supple-
mentary feeding affects red deer habitat selection. RSFs are statistical
models to assess habitat selection, contrasting resource use by animals
with resource availability (Boyce and McDonald, 1999; Lele et al.,
2013). Relocation data typically represents resource use, whereas re-
source availability is sampled using random locations distributed across
a predefined range. We sampled use and availability in a 1:1 ratio
within the 95% local convex hull nonparametric kernel home range
(Getz and Wilmers, 2004; Getz et al., 2007) of each GPS collared in-
dividual. First, we assessed seasonal differences in habitat selection of
red deer (H1). Therefore, we established a set of 13 candidate models
based on all or specific combinations of landscape variables as fixed
effects (Supplementary Table S1).
We included habitat types as dummy variables, and chose to exclude
‘other non-forest land’ to avoid rank deficiency (when all dummies add
up to 1) in the models. We included the interaction term ‘season’ on all
or on specific fixed effects to assess how habitat selection changes
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Forest Ecology and Management 407 (2018) 166–173
seasonally and according to specific landscape variables. We used the
Akaike Information Criterion corrected for small sample sizes (AICc)
and its difference values (ΔAICc) and model weights (w) for model
selection. We considered the least complex model within a 0–2 ΔAICc
range as the most parsimonious model (Burnham et al., 2011). We
quantified the relative importance of seasonal change in resource se-
lection specifically for each landscape variable included in the most
parsimonious model. Therefore, we systematically in- and excluded the
interaction term ‘season’ on a specific fixed effect, refitted the model,
and calculated the change in AICc (term specific ΔAICc, hereafter ‘Δc’).
Second, we assessed red deer habitat selection during the supplemen-
tary feeding season, and how distance to feeding stations mediates se-
lection for browsing and bark-stripping vulnerable habitat (H2). We
constructed a set of 8 candidate models based on the model structure of
most parsimonious model to assess H1, and allowed the ‘distance to the
nearest feeding station’ interact with browsing or bark-stripping vul-
nerable habitat types (Supplementary Table S2).
We included ‘individual deer’ as a random effect on the intercept in
all candidate models to account for sample size differences between
deer and spatial dependencies in the data. We decided to ignore ‘year of
monitoring’ as a random effect in the models due to the low number of
factor levels (3 levels, whereas minimum 5 are advised for inclusions as
a random effect), or as a fixed effect to reduce model complexity (Zuur
et al., 2009). We fitted the RSFs with logistic mixed-effect regression
models in R (R Development Core Team, 2013) with the ‘lme4’ package
(Bates and Maechler, 2010), and used the ‘MuMIn’ package for model
selection (Barton, 2012). We assessed collinearity among continuous
variables with a Pearson correlation test and considered correla-
tions > 0.6 as collinearities. We visually assessed data dependencies
between continuous variables and habitat types with conditional box-
plots and did not include non-independent variables in the same can-
didate model. We used actograms for red deer occurrence by habitat
types to enhance our understanding on diurnal and seasonal patterns in
red deer habitat use.
3. Results
We obtained 29799 valid GPS relocations of 11 deer (7 males, 4
females) between 2009 and 2011 (25 deer years, i.e., a year during
which an individual deer was monitored). We did not detect colli-
nearity among continuous variables. Altitude and habitat type, how-
ever, were not independent (Supplementary Fig. S1). Since habitat type
was our focal variable, we did not consider altitude in our candidate
models.
The most parsimonious model to assess seasonal differences in red
deer habitat selection contained all landscape variables with the in-
teraction term ‘season’, except for distance to roads (model weight
w = 0.728, Supplementary Table S1). We considered all other candi-
date models as inconclusive. By far the most important landscape
variable that distinguished red deer behavior during winter and
summer was distance to feeding stations: red deer strongly selected for
areas close to supplementary feeding stations during winter but not
summer, removing the seasonal interaction increased the model AICc
score with 5556.16 (Δc, Fig. 1).
Lesser pronounced seasonal shift in habitat selection behavior were
also apparent in relation to (in order of relevance) bark-stripping vul-
nerable forest (Δc = 313.02), slope steepness (Δc = 141.63), not vul-
nerable forest (Δc = 129.55), alpine meadows (Δc = 122.3), young
forest (Δc = 113.16), and windthrow areas (Δc = 97.01). During
summer, red deer selected for windthrow areas, young forests, mea-
dows, and alpine meadows and showed no clear preference or avoid-
ance for not vulnerable forest and bark-stripping vulnerable forests
(Supplementary Table S3). During summer, they also selected for
steeper slopes and areas relatively close to roads and feeding stations
(Supplementary Table S3). During winter, red deer avoided all land-
cover types except for meadows, which were strongly selected for, and
J.M. Arnold et al. Forest Ecology and Management 407 (2018) 166–173

Fig. 1. Kernel density distributions of GPS relocations (dark grey) of red deer and an equal number of randomly distributed points (light grey) within each deer’s home range during
winter (left) and summer (right). Red deer were monitored between 2009 and 2011 in a subalpine mountain forest in Austria.

they behaved similar towards roads as during summer (Supplementary 4. Discussion

Table S3).
The most parsimonious model to assess red deer habitat selection Our research provided two key findings. First, diversionary feeding
during winter contained an interaction between distance to feeding was the most important factor to explain red deer habitat selection
stations on young forest, windthrow areas, and bark-stripping vulner- differences between summer and winter in our study area (H1). Second,
able sites (Table 1). We discarded all other candidates, as they had diversionary feeding appears to release habitat selection pressure on for
ΔAICc values ≥96.6 (Supplementary Table S2). forestry vulnerable forest stands (H2), but this release decays with in-
Young forest, windthrow areas, and bark-stripping vulnerable for- creasing distance from the nearest feeding station.
ests were avoided compared to all other habitat types (Table 1), but Variation in food availability- natural or artificial- across seasons
only until a certain distances from supplementary feeding stations influences behavior, movement and habitat section in numerous wild-
(∼1.3–1.5 km, Fig. 2). Beyond these distances, young forest, wind- life species (Krebs et al., 1993; Hertel et al., 2016). For example, mi-
throw areas, and bark-stripping vulnerable forests were significantly gratory red deer appear to follow a phenological wave of vegetation
stronger selected for when compared to all other habitat types (Fig. 2). green-up across the landscape to move from summer to winter ranges
(Bischof et al., 2012); or, seasonal pulses of salmon (Oncorhynchus spp.)
Table 1 availability mediate movements and habitat selection of brown bears
Parameter estimates (β), standard errors (se), z and p-values of model terms retained in (Ursus arctos) that forage on this resource (Deacy et al., 2016). Variation
the most parsimonious model to assess red deer habitat selection during winter in a in food availability can be an important factor affecting life-history
subalpine managed forest in Austria in 2009–2011.
parameters in wildlife (Abrams, 1991), and is even a key factor in the
Model term β se z p-value evolution of mating systems (Emlen and Oring, 1977). Most resource
pulses occur naturally. However, anthropogenic food supplies become
Intercept 2.755 0.123 22.46 < 0.001 increasingly available for wildlife, for example, as crops, garbage and
Meadow 2.001 0.305 6.57 < 0.001
food waste, fishery discards, or through supplementary feeding (Mateo-
Alpine meadow −0.864 0.097 −8.86 < 0.001
Distance to feeding stations −2.811 0.041 −68.98 < 0.001 Tomás et al., 2015). As with natural foods, anthropogenic food supplies
Young forest −2.071 0.164 −12.62 < 0.001 can affect wildlife behavior, community assemblages, life history, social
Windthrow area −2.827 0.121 −23.41 < 0.001 organization, and mating systems (Craighead et al., 1995; Sahlsten
Bark-stripping vulnerable forest −3.013 0.105 −28.81 < 0.001 et al., 2010; Steyaert et al., 2014; Mateo-Tomás et al., 2015). Our re-
Not vulnerable forest −0.653 0.056 −11.74 < 0.001
Distance to roads −0.468 0.145 −3.22 0.001
sults add to the growing body of evidence that anthropogenic derived
Slope −0.002 0.001 −1.99 0.047 foods affect wildlife, as we show that supplementary winter feeding of
Distance to feeding stations · Young forest 1.514 0.132 11.45 < 0.001 red deer was the foremost important factor that steered red deer habitat
Distance to feeding stations · Windthrow 2.198 0.096 23.01 < 0.001 selection in our study area.
area
In our study area, winter feeding stations released red deer habitat
Distance to feeding stations · Bark- 1.828 0.077 23.70 < 0.001
stripping vulnerable forest selection pressure on vulnerable forest stands between 1.3 and 1.5
kilometers. In this ‘zone of release’, red deer selection for vulnerable
stands is lowered during winter, which may also reduce browsing and
bark-stripping. Despite the intensive feeding regime, however, deer
Red deer use of feeding stations increases over feeding time (from damage by browsing and bark-stripping is exerted and can locally even
October to May), and used feeding station throughout the day towards reach critical levels for regeneration forests and for timber production
the end of feeding season. With the end of feeding season, start of ve- (Gerhardt and Hochbichler, 2011). Several mechanisms may explain
getation season and calving period they abruptly started using young the persistently high levels of deer damage in our study area. First, the
forests and wind throw areas, and started to select strongly for thickets spatial distribution of the feeding station may not be sufficiently dense
and pole wood stands during daytime (Fig. 3). to cover the entire area with zones of release. Hence, a substantial
proportion (62%) on of the study area is not released from red deer

169
J.M. Arnold et al. Forest Ecology and Management 407 (2018) 166–173

Fig. 2. Predicted effects of distance to the nearest feeding stations on red deer selection for habitat types vulnerable for deer-induced damages during winter. For visualization purposes,
we refitted the most parsimonious mixed effect logistic regression model without random intercept using a generalized linear model.

habitat selection in vulnerable stands and might thus exert high damage
levels. In addition, red deer population densities in the immediate vi-
cinity of feeding stations can be very dense, which may result in severe
local damages (Jerina, personal communication). Second, reduced ha-
bitat selection pressure does not necessarily implies lowered damage.
Despite using these stands to lesser extent, red deer may still incur se-
vere damage. Third, red deer can cause severe damage in vulnerable
forest stands during summer as well, even if or because of natural
forage is abundant and attractive in those stands (Gill, 1992). Fourth,
the current hunting management appears to be unable to control deer
impacts on the forest to a satisfactory level. One must be aware that
feeding ungulates leads to the premise of a higher effort in controlling
deer numbers (Deutz et al., 2009). To a certain extent, such satisfactory
levels could be reached by reducing deer densities through increasing
hunting quotas and spatiotemporal adjustment of hunting effort, al-
though deer can adjust their behavior with increasing age by learning
from negative experience (such as drive hunts), and pass on this
strategy to other animals of the group (Thurfjell et al., 2017). To reduce
damage in vulnerable stands, we suggest that hunting effort could be
focused on vulnerable forest stands during periods of high risk for da-
mage (Beguin et al., 2016) with sparing feeding areas of low suscept-
ibility such as meadows or wind throw areas not cleaned from debris. In
addition, hunting pressure should remain high for prolonged periods of
time (e.g. several years) (Simard et al., 2008) and focus on female red
deer (Milner et al., 2006).
Supplementary feeding may be in conflict with forestry goals in the
long term. It artificially increases natural carrying capacity, especially
of winter habitat, and animal population densities (Putman and Staines,
2004; Peek et al., 2002). Increased deer population densities may
propagate into increased browsing or bark-stripping damage, and blur
the effectiveness of diversionary feeding. Seizing supplementary
feeding could reduce red deer population densities, as for example in
the Yellowstone grizzly bear population (Craighead et al. 1995), which
may in turn reduce forest damage.

Fig. 3. Actograms of red deer activity within 125 m of a winter feeding station, or within habitat types vulnerable for red deer browsing (‘Young forest’, ‘Windthrow’) or bark-stripping
(‘Bark-Stripping’). Each point represents the frequency of occurrence of GPS marked red deer (N = 11) within a specific habitat type at a specific time on the day and day of year
(2009–2011). The size of the points reflects the absolute number of GPS relocations at a specific time divided by a scaling factor (i.e., factor 10 for ‘Feeding stations’ and factor 5 for the
other habitat types). We introduced a 3-hour noise jitter on ‘Time of day’ for visualization purposes. The dotted lines represent sunrise and sunset. Note that winter feeding occurs between
October (‘Day of year’ ∼300) and May (‘Day of year’ ∼ 120).

170
J.M. Arnold et al.
Supplementary feeding can have a myriad of indirect and un-
expected effects on the target species (Putman and Staines, 2004;
Milner et al., 2014). Risks associated with feeding (as reviewed in
Putman and Staines, 2004) include increased competition among in-
dividuals (Wiersema, 1974; Linn, 1986; Schmidt, 1992; Seivwright,
1996) and imbalanced diets (Doman and Rasmussen, 1944; Wiersema,
1974). Aggregating wildlife around feeding stations is also associated
with the potential for increased parasites and disease transmission
(Putman and Staines, 2004). For example, supplementary feeding is
generally acknowledged to foster the prevalence and maintenance of
bovine tuberculosis (caused by Mycobacterium bovis or M. caprae) in
wildlife populations by animals aggregating around feeding stations
(Castillo et al., 2011; Schöning et al., 2013, Fink et al., 2015). Fur-
thermore, supplementary feeding can reduce natural selection potential
in juveniles (Schmidt and Hoi, 2002), and it can increase population
densities, thereby delaying calving dates (Coulson et al., 2003; Walling
et al., 2011), and reduced calf birth weight (Stopher et al., 2014) and
survival (Coulson et al., 1997; Walling et al., 2011). Increased hind
population densities typically delay age of hinds’ first reproduction
(Kruuk et al., 1999a) and decline fecundity (Kruuk et al., 1999b).
Supplementary feeding can furthermore affect learning and adaption,
with the potential risk on reliance on supplementary foods (Putman and
Staines, 2004), and loosing natural overwintering and migratory stra-
tegies (Schmidt, 1993; Luccarini et al., 2006). In addition to unexpected
effects on the focal wildlife species, supplementary feeding, can also
have unexpected effects on non-target species and throughout the
ecosystem (Robb et al., 2008; Milner et al., 2014; Selva et al., 2014).
For example, it can alter nutrient redistribution (Robb et al., 2008) and
modify behavior of non-target species (Selva et al., 2014), provoke
shifts in vegetation composition and diversity (Rinella et al., 2012),
inhibition of natural processes such as endozoochorous seed dispersal
(Steyaert et al., 2009), and increase of depredation risk of ground
nesting birds (Oja et al., 2015).
Diversionary feeding is the most commonly applied protective
measure to manage forests for deer damage (Putman and Staines, 2004)
although many studies have shown that this measure is counter-
productive and even mediating deer damage (Náhlik, 1995; Luccarini,
2006). However, several alternative measures can be successfully ap-
plied. Implementing specific silviculturals methods (e.g., thinning,
clear-cut regime) can steer red deer habitat selection. Such measures
can balance deer numbers and settling stimulus (i.e., attractiveness of a
certain area in terms of plant quantity and quality, thermal conditions
or cover characteristics) with forage supply, with a focus on the winter
carrying capacity. Enhancing natural food availability likely lowers
deer impact susceptibility (Reimoser et al. 2006; Jarnemo et al., 2014).
At forest stand scale, silvicultural actions can impact forage resources in
terms of quantity, quality, and distribution (Beguin et al., 2016). Sil-
viculture directly affects environmental conditions for deer through its
effects on stand cover, understory vegetation, tree regeneration, stand
structure, and forest edge properties (Wagner et al., 2011). For ex-
ample, natural regeneration after selective harvesting of single trees or
group felling can provide a valuable forage resource for deer, and en-
hances patchiness of forage availability over larger areas. It could also
make an area less attractive, as it would increase search effort to find
such patches. Such a harvest regime would also introduce more blurred
edges to the forest, rendering it less attractive for deer (Reimoser and
Gossow, 1996; Storms et al., 2006). Further, early thinning to improve
light penetration to the ground and enhance natural regeneration of
forage plants (Kramer et al., 2006) or tree seedlings (Armstrong et al.,
2003). Other methods, such as progressive clear-strip systems or shel-
terwood cuts (Reimoser et al., 2006), or fostering advanced tree re-
generation (Kuiters et al., 1996; Partl et al., 2002) could be applied to
improve the balance of forage-independent settling stimulus and food
availability (Reimoser et al., 2006) by improving landscape hetero-
geneity and consequently lowering damage susceptibility. In general,
silvicultural means could be used to aim for a decreased attractiveness
171
Forest Ecology and Management 407 (2018) 166–173
of the habitat while at the same time increasing the forage supply
(Gerhardt et al., 2013), and forests’ resilience against environmental
factors in general.
Trees can also physically be protected from deer by fences, tree
guards, and natural protection, such as brushwood hedging (Armstrong
et al., 2003). For example, dead branches or patches of shrubs and
bramble can protect young, naturally regenerated trees from deer
browsing (Sanderson, 1996; Hamard and Ballon, 1998; Kupferschmid
and Bugmann, 2005; Rammig et al., 2007). This is also true for wind-
throw areas: by not cleaning areas from fallen trees and branches, tree
regeneration can be protected from browsing by coarse woody debris
(Nagel et al., 2006; Peterson and Pickett, 1995). Several studies have
shown that ungulates are not causing significant damage to natural tree
regeneration in (uncleared) windthrow gaps (Senn et al., 2002; Angst
et al., 2004; Nagel et al., 2006), which in production forests are usually
cleaned from woody debris, as plant biomass increases very rapidly and
to exceed their consumption needs and browsing capacity (Senn et al.,
2002). By using chemical repellents on trees (mainly apical buds), deer
browsing on regenerating seedlings and trees can be reduced (Nolte,
1998).
During winter metabolic rates, activity and food intake in red deer
strongly decrease (Arnold et al., 2004; Turbill et al., 2011). This winter
adaption can be annulled by disturbing animals, consequent higher
needs for food intake and higher activity levels may provoke damages,
leading up to 30% of higher damage rates (Arnold, 2016). Therefore,
disturbances should be avoided from winter solstice when metabolic
changes begin to late winter.
5. Conclusion
One of the foremost important goals of supplementary feeding of
red deer is the prevention of damage on production forests, while
maintaining high deer densities for hunting purposes. Our results pro-
vide evidence that diversionary feeding can be effective to spatio-
temporally redistribute a red deer population to reduce deer impact on
vulnerable stands in intensively managed production forests. Despite
intensive winter feeding and hunting, high levels of browsing in re-
generating forest stand and bark-stripping persist. Several factors may
explain this, and include the spatial distribution and density of the
feedings stations, animal behavior, and the inherent conflict between
supplementary feeding for forestry (minimizing red deer impact and
potentially increasing red deer population density) and hunting pur-
poses (controlling for red deer number and concurrently sustaining
higher number of red deer). In addition, numerous other factors need to
be considered to control deer impact and to balance specific forestry
and wildlife management goals. Hence, a holistic and science based
management to achieve such goal is required, and includes silvicultural
measures (e.g. to enhance natural food availability), spatial planning of
land use (e.g. feeding stations, wildlife refuge zones), and stakeholder
involvement (e.g. hunting, forestry, tourism). Our understanding of the
ecological effects and conservation implications of supplementary
feeding wildlife is still very limited (Robb et al., 2008; Mateo-Tomás
et al., 2015), including the interplay numerous of biotic and abiotic
factors that affect deer-induced forest damage (Gerhardt et al., 2013).
Besides effectiveness, also potential risks and undesired effects of sup-
plementary feeding should be taken into account when applying such
strategies. We argue that landowners, managers, and conservationists
carefully consider such effects when implementing supplementary
feeding strategies.
Acknowledgements
This work was supported by the Austrian Research Promotion
Agency (FFG, BRIDGE-research grant, project number 820112), Vienna,
Austria, and the Princely Schwarzenberg Family Foundation, Vaduz,
Liechtenstein. We thank the Princely Schwarzenberg Family
J.M. Arnold et al. Forest Ecology and Management 407 (2018) 166–173

Foundation and all employees of its forest enterprise Murau, especially Science 197, 215–223.
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Flachmeier, Christoph Gottsbacher, Thomas Reiter, Bernd Scacel, Fink, M., Schleicher, C., Gonano, M., Prodinger, W.M., Pacciarini, M., Glawischnig, W.,
Gebhard Auer, Herbert Gerhardter, and Michael Stremminger. Ryser-Degiorgis, M.-P., Walzer, C., Stalder, G.L., Lombardo, D., Schobesberger, H.,
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(Steirische Landesjägerschaft, Kärntner Landesjägerschaft and Gerhardt, P., Hochbichler, E., 2011. Waldbauliche Kennzahlen für die
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University of Natural Resources and Life Sciences, Vienna.
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