Habitat use, bed-site selection and mortality rate in
neonate fallow deer Dama dama
Authors: Kjellander, Petter, Svartholm, Ida, Bergvall, Ulrika A., and
Jarnemo, Anders
Source: Wildlife Biology, 18(3) : 280-291
Published By: Nordic Board for Wildlife Research
URL: https://doi.org/10.2981/10-093
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Original article
Wildl. Biol. 18: 280-291 (2012)
DOI: 10.2981/10-093
Ó Wildlife Biology, NKV
www.wildlifebiology.com
Habitat use, bed-site selection and mortality rate in neonate fallow
deer Dama dama
Petter Kjellander, Ida Svartholm, Ulrika A. Bergvall & Anders Jarnemo
An understanding of mortality patterns, and especially the variation in juvenile mortality, is an important component in
vertebrate population dynamics. Our study investigates, for the first time, neonate mortality and two levels of spatial
behaviour, in a free-ranging fallow deer Dama dama population in southwestern Sweden. In the summers of 2008 and
2009, 36 fawns were marked with radio-collars. Neonate mortality calculated by the Kaplan-Meier method was 23.6%.
Mortality caused by predation was low, since only one of eight non-surviving fawns died from predation, probably by red
fox Vulpes vulpes. The spatial behaviour of the neonates was examined by habitat selection at home-range level, which in
fact is a selection made by the mother, and at bed-site level within that habitat. Compositional analysis revealed a
significant preference for arable land, pasture and coniferous forest between 5-15 m high, compared to young forest.
Selected bed sites showed significantly lower visibility and higher amount of canopy cover than random sites. Surprisingly,
we did not find any relationship between canopy cover and visibility in selected bed sites while it showed a significant and
negative relationship at random bed sites. We interpret this finding as while high canopy cover and low visibility covary at
the habitat level, fawns seem to select these two bed-site variables independently, perhaps for thermoregulatory reasons.
Since there are few predators in our study area and predation pressure is low, this behaviour is not connected to actual
survival rates in this area, but would rather be in support of the hypothesis of ’pleiotropy’ as thermoregulatory reasons for
bed-site selection in neonate fawns might be the most important contemporary selection force in the absence of large
predators.
Key words: anti-predatory behaviour, bed sites, Dama dama, fallow deer, habitat selection, neonate mortality, predation,
thermoregulation
Petter Kjellander, Ida Svartholm & Anders Jarnemo, Grimsö Wildlife Research Station, Department of Ecology, Swedish
University of Agricultural Sciences, SE-730 91, Riddarhyttan, Sweden - e-mail addresses: Petter.Kjellander@slu.se (Petter
Kjellander); Ida.Svartholm@slu.se (Ida Svartholm); Anders.Jarnemo@slu.se (Anders Jarnemo)
Ulrika A. Bergvall, Department of Psychology, School of Philosophy Psychology and Language Sciences, University of
Edinburgh, 7 George Square, EH8 9JZ Edinburgh UK, and Grimsö Wildlife Research Station, Department of Ecology,
Swedish University of Agricultural Sciences, SE-730 91, Riddarhyttan, Sweden - e-mail: Ulrika.Alm.Bergvall@slu.se
Corresponding author: Petter Kjellander
Received 20 August 2010, accepted 31 March 2012
Associate Editor: Stefano Focardi
An understanding of mortality patterns, and especially the variation in juvenile mortality, is an
important component in vertebrate population dynamics (Gaillard et al. 1993, Aanes & Andersen 1996,
Van Moorter et al. 2009). Ungulate species generally
show a U-shaped mortality curve with the highest
mortality occurring at early and late periods of life
(Caughley 1966), and the rate of neonate mortality in
temperate ungulates sometimes exceeds 50% (Lin280
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nell et al. 1995, Aanes & Andersen 1996, Jarnemo et
al. 2004). This first summer mortality can be caused
by disease, starvation, hypothermia, parasites and
accidents, but in areas where predators are present,
the number one mortality factor for neonate ungulates is predation (Linnell et al. 1995).
Due to the high risk of predation, neonate
ungulates and their mothers have evolved a range
of anti-predator strategies (Lent 1974, Ciuti et al.
Ó WILDLIFE BIOLOGY 18:3 (2012)
2006, Bongi et al. 2008). Based on the type of motherinfant relationship, most ungulate species can be
categorised as either a ’hider’- or a ’follower’-type
(Lent 1974). In species categorised as hiders, the
mother and the infant stay separated and out of
contact for long periods of time, during which the
infant stays hidden in the vegetation. Most species of
cervids fall into this category even though the hider
phase can vary in time from 2-3 days in some species
(e.g. Siberian ibex Capra ibex, mouflon Ovis orientalis and red deer Cervus elaphus), to 2-4 months in
others (e.g. Uganda kob Kobus kob and reedbuck
Redunca redunca; Lent 1974). In follower species,
newborns actively follow the mother and maintain a
close and frequent contact from the start. Many of
the follower species are seasonal movers, associated
to grassland or tundra habitats (e.g. the caribou
Rangifer tarandus and the muskox Ovibos moschatus;
Lent 1974, 1991). This strategy provides protection
against predators by allowing the mother and fawn
to remain in a group, relying on group defense and
permitting extensive movements (Lent 1991). In case
of relaxed selection because of the loss of one or more
predator species, anti-predator behaviour like these
can still persist, which is referred to as ’the ghost of
predators past’ hypothesis (Byers 1997a, Blumstein
et al. 2006).
The fallow deer Dama dama is classified as a hiderspecies, since the fawns lie hidden during the first two
weeks of their life (San José & Braza 1992, Ciuti et al.
2006). The mother typically stays within hearing
distance of her fawn and visits several times per day
to feed and groom it (Chapman & Chapman 1997)
and fallow deer females rarely move further away
than 400 m from the fawn (U.A. Bergvall, unpubl.
data). Since a secluded bed site is the main protection
for these fawns, bed sites could be expected to be
selected thoroughly in order to provide good cover.
Although the change of hiding place is typically
initiated by the female, the exact position of the
actual bed site is chosen by the fawn (Lent 1974,
Heugel et al. 1986). Bed-site selection by fawns of
other hider-type species (e.g. roe deer Capreolus
capreolus (Linnell et al. 1999) and white-tailed deer
Odocolius virginianus (Heugel et al. 1986)) has been
examined previously. However, this behaviour has
not yet been studied in fallow deer.
As well as serving as a protection against predators, the use of cover and the importance of bed-site
selection can also be expected to be affected by
climatic factors (Heugel et al. 1986, Mysterud &
Østbye 1999). Ungulate neonates are vulnerable to
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cold and wet weather, and hypothermia is often
mentioned as the main cause of non-predator mortality in the review of Linnell et al. (1995; see also
Olson et al. 2005). The high neonate mortality
recorded for fallow deer kept in open paddock
conditions on deer farms is thought to be a result of
the lack of shelter from high rainfalls (Putman et al.
1996). Thus, at a higher level, the selection of main
habitat types might be very important, since different
habitats may provide different opportunities for the
fawns to choose a protected bed site. Selection is
therefore expected to also act towards fawns choosing the most beneficial bed site also for thermoregulatory reasons, for a given weather condition (Van
Moorter et al. 2009). Beside selection for cover
against predators, bed-site selection in neonate
fallow deer might, therefore, also be an example of
Byers (1997a) ’pleiotropy’, i.e. an anti-predatory
behaviour that remains because it also has an
alternative function (see also Stearns 2010).
There are obviously many factors affecting habitat
selection among herbivores, such as food quality and
abundance, availability of shelter and cover from
predators or unfavourable weather conditions, but
also social structure and population densities (Putman 1988, Sinclair et al. 2006). Furthermore, the
presence of fawns leads to changes in social behaviour and space-use of calving females (Shackleton &
Haywood 1985, Schwede et al. 1993, Bon et al. 1995,
Kohlmann et al. 1995, Tufto et al. 1996, Ciuti et al.
2006, Bongi et al. 2008). Ciuti et al. (2006) showed
that female fallow deer in Italy adopt an antipredator behaviour in their habitat selection, since
calving females to a higher extent used suboptimal
habitats that offered the best cover and thereby
reducing the predation risk for their fawns, compromising their own energy intake. In order to be close to
their fawns, calving females also reduced the size of
their home ranges, using areas between two or three
times smaller than the areas used by non-calving
females (Ciuti et al. 2006).
Despite the fact that the fallow deer is one of the
most widely distributed species of deer in the world
(Chapman & Chapman 1997), relatively little research has been done regarding the ecology of the
species (Borkowski & Pudelko 2007), and especially
on free-ranging fallow deer in Northern Europe and
Scandinavia (Chapman & Chapman 1997, Carlström & Nyman 2005). The species was first introduced to Sweden in the 1570s, and it is now welldistributed in the form of scattered occurrences over
the southern part of the country (Carlström &
281
Nyman 2005). The reported annual harvest of fallow
deer in the wild has increased from 1,000 animals in
1955 to almost 17,000 in 2005 (Bergström & Danell
2009). With an increasing population, the need of
good management plans increases, and an important
part of population dynamics is the understanding of
mortality patterns. Preliminary and indirect data,
based on harvested females, indicate that approximately 30% of neonate fallow deer in a population at
Koberg in southwestern Sweden do not survive their
first summer (P. Kjellander, unpubl. data). Our study
is the first trying to verify these indicative findings
and to investigate causes of neonate mortality.
The aims of our study were therefore to investigate
1) the level of neonate mortality in a free-ranging
fallow deer population in Sweden, 2) the habitat
selection, made by the mother, and 3) the environmental factors affecting bed-site selection by the fawn
that may influence neonate survival. Because of the
anti-predatory origin of the hider strategy and the
sensitivity to thermal conditions seen in ungulate
neonates, we expected the fallow deer fawns to select
bed sites that provided more cover and better
concealment than random sites (Lent 1974, Putman
et al. 1996, Linnell et al. 1999). As a consequence, we
also predicted bed-site selection to affect fawn
survival, e.g. we expected the surviving fallow deer
fawns to have chosen bed sites with better cover and
concealment compared to the non-surviving fawns.
Material and methods
year by a Distance Sampling procedure (Buckland et
al. 2001) to 17.7 and 19.9 animals/km2 in 2008 and
2009, respectively (P. Kjellander, unpubl. data.).
Other ungulates occurring in the area regularly are
roe deer, moose Alces alces and wild boar Sus scrofa
and occasionally red deer. Controlled hunting is
performed each fall (September-February, with a
pause during the rut), and potential predators beside
wild boar present in the area are the red fox Vulpes
vulpes and occasional visits of lynx Lynx lynx and
wolf Canis lupus.
Temperature and precipitation in our study area
was recorded by the Swedish Meteorological and
Hydrological Institute (SMHI) at the Gendalen
meteorological station, situated 12 km from our
study area. Mean monthly temperatures in our study
area during summer (June-August) usually range
between 15 and 178C, with the highest temperatures
during July (SMHI 2009). Mean monthly precipitation (June-August) range between 86 and 130 mm
with June and July normally being equally dry
whereas August has the highest rainfall (SMHI
2009). The two summers included in our study (2008
and 2009) did not diverge substantially from the
monthly mean weather conditions reported here, as
mean temperature ranged between 15.6 and 15.88C in
the two years, while there was slightly less rain fall
than average in June and July 2008 (60-64 mm) as
well as June 2009 (65 mm), while August 2008 and
July 2009 was unusually wet with 50-100% more rain
than the monthly average (197 and 162 mm, respectively).
Study area
Our study was performed at the Koberg estate (588N,
128E) in southwestern Sweden (Västra Götaland
County). Our study area (54.35 km2) is mostly
covered with different types of forest (79%), and the
remaining area consists of arable land and pastures
(16%), mires and marshes (2%), lakes, ponds, parks
and properties around houses (3%). The two most
common habitat types are coniferous forest . 15 m
(29%) and coniferous forest 5-15 m (15%; Winsa
2008). The open landscape at Koberg, consisting of
arable land and pastures, are today, to a large extent,
cultivated in order to improve wildlife habitats, and
supplementary food is also given during wintertime.
Free-ranging fallow deer has been present in the area
since the release of a few animals (approximately 20)
from an enclosure in the end of the 1920s (N.
Silfverschiöld, unpubl. data). The fallow deer population in our study area was estimated in April each
Data collection
Data was collected during the summers of 2008 and
2009. Fallow deer fawns were found, by either
searching the near surroundings of solitary females
observed standing by their own outside a group, or
by waiting for them to visit their hidden fawns.
Fawns could also be detected by chance when
searching probable fawning areas. The fawns were
caught during the first days in life, when they still
adopt a prone position, either by hand or with
landing nets. Sex and body mass of the fawns were
determined at capture together with other measurements such as length of metatarsus, heart girth and
body temperature. The fawns were ear-tagged with
coloured plastic tags and equipped with a radiotransmitter attached to an expandable collar with a
drop-off function. In 2008, 15 fawns were caught; 12
were fitted with VHF transmitters (Televilt, Lindesberg, Sweden) and only three were ear-tagged. In
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Ó WILDLIFE BIOLOGY 18:3 (2012)
2009, 24 fawns were caught, and this year, we marked
them with either a traditional VHF (N ¼ 21) or a
VHF/GPS transmitter (N ¼ 3; Telemetry solutions,
Concord, California, USA). GPS transmitters were
programmed to take one position every hour. All
transmitters were equipped with a mortality function
(motion-sensitive). Fawn age at capture was estimated based on a number of characteristics, such as the
texture of the fur, the status of the umbilical cord,
hoof abrasion and the behavioural reactions of the
fawn during capture, handling and release (Galli et
al. 2008). All possible sorts of concern to prevent
negative effects of the first capture was taken (e.g.
keeping silent and calm during handling, minimising
disturbance time (, 10 minutes) and number of
people at capture sites ( 2) and by rubbing grass on
hands and tools).
Fawn survival was normally checked daily for up
to four weeks of age, until mid July, then once a week
until mid September. Fawns marked with VHF
transmitters were located using three element yagi
antennas and receivers. The exact locations of the
fawns were estimated with a script built on the R
platform (R version 2.10.0, R development core team
(2006-2009)) by using bearings from three or four
reference points. If a fawn was observed during radio
tracking (N¼ 5 locations), the observed position was
used. In order to establish and categorise the cause of
death, the location of the found collar or the dead
fawn was investigated for markings or other signs of
predation, and the dead fawns were necropsied and
investigated for bite marks and signs of diseases.
Data on bed-site selection was collected from 21
June to 14 July 2009 for fawns marked during the
same period. Each fawn was radio-tracked and
carefully approached in order to localise their exact
bed site. The location of the bed site was noted with a
hand-held GPS, marked with a plastic strip and
examined the following day(s) when the fawn had
moved to another location. For each bed site, a
random bed site was selected using a random table
where direction and distance from the actual bed site
was given. An inflated basket ball, representing the
fawn, was placed in the bed site and approached from
each of the four cardinal directions, by the same
person throughout the whole study. We started
measuring from a maximum distance of 50 m, and
then we recorded the distance from which the ball
was first detected at a height of 50 cm above the
ground. Canopy cover above the bed site was
estimated in percent from a height of approximately
1.5 m and a short description of the bed site was
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made. The same procedure was repeated for the
randomly selected bed sites. In order to make a
correct comparison between the selected and the
randomly chosen bed sites, they had to be situated in
the same habitat type or forest stand. In case the
random distance reached out of the current habitat,
the remaining distance were measured back from the
border of the habitat towards the selected bed site.
Data analysis
As the weather conditions and population density
estimates were rather constant between years and
annual sample sizes fairly small, we pooled data over
the two years. Following that, we estimated fawn
mortality by the Kaplan-Meier method (Pollock et
al. 1989) for the first 70 days of age on data from both
years. This method allows for animals to enter the
study at different times, since the estimated birth date
is set as day 1 for all fawns. We tested differences in
survival rates between male and female fawns with
the G-test of goodness of fit.
The marking positions of each fawn were not
included in the habitat-use analysis, since most of the
fawns were found in open areas, due to a higher level
of visibility in such habitats. We analysed the data on
fawn positions in Arc GIS 9.3 and used a satellite
generated digitalised map to define each habitat.
The map was derived and developed from the
Swedish CORINE Land Cover map to ’Svensk
marktäckedata’ (SMD) with a 25 3 25 m pixel
resolution (Lantmäteriet 2004). We randomly moved
fawn positions that ended up between two habitats
into one of the two neighbouring habitats (five out of
726 positions). The habitat type ’lakes and ponds,
open water surface’ was not considered available
habitat and therefore removed from further analysis.
We accordingly based data on home ranges and
habitat use on locations collected during the
estimated first month of each fawn’s life, starting
with the first marked fawn 21 June and ending 15
August with the latest marked fawn (marked 14
July). We calculated home ranges with the minimum convex polygon method (MCP) using the
extension Hawth’s Tools. Random points were
then generated within each home range in order to
estimate the habitat availability for each fawn. We
analysed the habitat use within the MCP home
range with compositional analysis (CA; Aebisher et
al. 1993) in the software Resource selection (Resource selection for Windows, version 1.00 Beta
8.1, 1999 Fred Laban). To reduce the number of
habitat types for the CA, we combined similar and
283
rare habitat categories as follows: land in active
agricultural management as arable land (ARABLE), open grasslands never ploughed but previously or currently grazed by live stock as pastures
(PASTU), new or recently replanted clear-felled
forests with sprouts up to 2 m height (CLEAR),
younger forest with a tree height between 2-5 m
(YOUNG), mixed conifer-deciduous forest . 5 m
(MIXED), pure coniferous forest of 5-15 m height
(CON 5), pure coniferous forest . 15 m (CON 15)
and finally wet areas as various types of mires and
over-grown water ponds (WETLAND). Since CAs
require no missing values for either habitat use or
habitat availability, we replaced missing values
with 0.001. Considering the small sample sizes
(number of fawns) and the eight habitat classes, we
arbitrarily discarded fawns with , 15 positions in
order to balance the data set somewhat for the CA
(Aebisher et al. 1993). Hence, only data from 14
fawns from both years could be used, since many of
the fawns died or lost their collars too early
resulting in a mean of 43 6 6.47 SE locations per
fawn (range: 15-81 locations). When we corrected
for the number of fixes as a covariate in a resource
selection function model (RSF) with a logistic
approach, it did not change our conclusions. We
therefore decided to only account for the result
from the CA. Because of the fact that fawns died
before we acquired enough positions, we were
unable to calculate home ranges and test for differences in habitat choices between surviving and
dead fawns. Even though we are referring to the
positions and habitat use of the fawns, this selection, as discussed earlier, is in fact made by the
fallow deer female.
We used the mean distance (of the four cardinal
distance measures), at which the model fawn was
detected, as a measure of visibility. Firstly, we tested
for differences in visibility between the selected bed
sites and random bed sites, between individual fawns,
fate (dead or alive) and the interaction between site
(selected and random) and fate (dead or alive) with
an analysis of variance (repeated ANOVA). Secondly, we analysed the data for possible differences in
visibility between the different habitats. Results are
presented as means with standard deviation, if not
otherwise specified. In order to analyse data on
canopy cover, which were recorded in percent,
pffiffiffiffiffiffiffiffian
ffiffiffi
arcsine-squareroot transformation (y ¼ sin-1 x)
was required (Krebs 1999). The transformed values
on canopy cover were then tested in the same way as
the data on visibility. The presented means (with SD)
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are nevertheless based on the original, non-transformed data. We tested differences in bed-site selection between surviving and non-surviving fawns with
repeated ANOVAs. All statistical tests, beside the
CA, were performed in StatView 5.0.1.
Ethical approval for our study was granted by the
Gothenburg Board for Laboratory Animals (Dnr:
187-08 & 405-08).
Results
A total of 39 fawns (18 males, 21 females) were
captured and marked during our study; 15 fawns in
2008 and 24 fawns in 2009 (Table 1). In 2008, three
fawns were only ear-tagged and they could therefore
not be included in the mortality calculations. Thus,
out of a total of 36 fawns (17 males, 19 females) eight
fawns died, of which five were males and three
females. The death of one fawn could likely be
assigned to red fox predation, since the collar was
found with fox bite marks, traces of blood and scent
of red fox urine. Six fawns died from starvation and
one of unknown causes. All of those six starved fawns
had no traces of milk in the gastro-intestinal tract,
implying that they had never been fed by their
mothers. The cumulative mortality according to the
Kaplan-Meier analysis was 23.6% (6 0.07 SE) for all
fawns. Even though not significantly different, the
mortality was 16.4% (6 0.08 SE) in females and
32.1% (6 0.1 SE) in males (G ¼ 0.27, df ¼ 1, P ¼ 0.6;
Fig. 1). The mean date of birth was 23 June (6 5.0
days) for the marked fawns.
Habitat use
According to the CA, the habitat use of the fawns,
selected by their mothers at the home-range scale
differed significantly from a random choice (v2 ¼
14.65, df ¼ 7, P , 0.05). The habitat classes arable
Table 1. Numbers, sex and mortality of neonate fallow deer fawns
marked at Koberg in southwestern Sweden in 2008 and 2009. In
2008, three of the 15 marked fawns were only ear-tagged (one male
and two females).
Number marked
Males
Females
Number dead
Males
Females
2008
2009
Both years
15
6
9
24
12
12
39
18
21
3
1
2
5
4
1
8
5
3
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Figure 1. The cumulative survival of fallow deer neonate females
(N ¼ 19, black line) and males (N ¼ 17, dashed line) at Koberg,
Sweden, during their first summer of life. Censored animals are
indicated by j, i.e. individuals lost to our study due to dropped
collars. Data from 2008 and 2009. The gender specific survival rate
indicated in the figure did not differ significantly from each other
(see the result section for more details).
land (ARABLE), pasture (PASTU) and coniferous
forest between 5-15 m (CON 5) were significantly
more preferred (t . 2.282, df ¼ 13, P 0.040 in all
cases) than young forest (YOUNG; Fig. 2), while no
significant differences in use of the remaining habitat
classes were found (CLEAR, MIXED, CON 15 and
WETLAND; t , 1.843, df ¼ 13, P . 0.088 in all
cases).
Bed-site selection
A total of 77 bed sites from 23 marked fawns in 2009,
1-5 bed sites for each fawn, was visited and measured
during our study (one fawn lost its collar during the
first 24 hours).
Out of 77 bed sites, 25 (32.5%) were situated in the
habitat class ARABLE, 19 (24.7%) in CON 15 and
12 (15.6%) in MIXED. YOUNG, CON 5 and
PASTU had the least number of bed sites (6.5% in
each habitat type). Six bed sites (7.8%) were situated
on CLEAR and no bed sites were found in WETLAND. Of all bed sites, 12 (15.6%) were also situated
in an edge zone, i.e. within 5 m of another habitat
type, mostly between forest and open field. Three bed
sites on ARABLE were situated within the edge of
some kind of forest. Eight bed sites in forest were
situated within the edge of ARABLE (four of the bed
sites in CON 5, one in CON 15 and three in MIXED).
One bed site on CLEAR was situated in the edge zone
of ARABLE.
ARABLE was the most used habitat for bed sites
by surviving fawns (30.3%), as well as for the fawns
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Figure 2. Habitat selection by fallow deer fawns (6 SD) at Koberg in
southwestern Sweden. Habitat classes to the left of the symbol . are
selected over those to the right. *¼P , 0.05 (compositional analysis)
indicate significant differences between those habitat classes
connected by a line. ARABLE ¼ arable land, PASTU ¼ pasture,
CON 5¼coniferous forest 5-15 m, CON 15¼coniferous forest .15
m, WETLAND ¼ wet areas, MIXED ¼ mixed forest . 5 m,
CLEAR ¼ clear felled areas, forest up to 2 m, YOUNG ¼ young
forest 2-5 m.
that did not survive (45.5%). This habitat had the
lowest canopy cover (4.8% at the selected bed sites),
but also a low visibility (3.1 m).
Visibility
There was a significant difference in visibility between
bed sites of the individual fawns (minimum: 1.4 6 0.5
m; maximum: 41.6 6 0 m; F22, 54 ¼ 4.15, P , 0.0001)
and the selected bed sites showed significantly lower
visibility (4.29 6 6.0 m) than the random bed sites
(8.07 6 8.3 m; F1, 22 ¼ 31.8, P , 0.0001). Visibility
between bed sites of surviving and non-surviving
fawns did not differ (F1, 21 ¼2.45, P , 0.133; Fig. 3A).
The visibility of the bed sites differed significantly
between habitat types (F6, 70 ¼ 3.28, P ¼ 0.007). Even
though CON 15 was the second most chosen habitat
category for bed sites, the visibility in this habitat,
based on random bed sites, were the highest with a
mean of 15.4 6 9.0 m (Fig. 4 and Table 2). However,
the selected bed sites in CON 15 had a mean visibility
of 6.9 6 4.5 m, which was lower than the mean
visibility for the selected bed sites in CON 5 (8.2 6 8.5
m). The habitat classes with the lowest visibility were
ARABLE (4.6 6 7.8 m), PASTU (4.2 6 3.0 m) and
YOUNG (4.3 6 1.6 m).
Canopy cover
There was a large difference in canopy cover, ranging
from 0 to 100%, between bed sites of the individual
285
Figure 3. Mean bed-site visibility (A) and
cover (B) of the selected (white) and random
bed sites (light grey) of 18 surviving and five
non-surviving fallow deer fawns (6 SD) at
Koberg in southwestern Sweden, in the
summer of 2009.
fawns, and the selected bed sites had a significantly
better mean canopy cover (24.8 6 32.3%) than the
random bed sites (5.3 6 17.3%; F1, 22 ¼ 36.2, P ,
0.0001; see Fig. 3B), but canopy cover between bed
sites of surviving and non-surviving fawns did not
differ (F1, 21 ¼ 0.06, P¼ 0.8; see Fig. 3B). The amount
of canopy cover differed significantly between habitats (F6, 70 ¼ 4.03, P ¼ 0.002; Fig. 5), and ARABLE,
PASTU and CLEAR had almost no canopy cover at
all at the random sites, with a mean cover , 1%
(ARABLE: 0.6 6 0.5%, PASTU: 0.6 6 0.5%,
CLEAR: 0.2 6 0.4%). However, the measurements
from the selected bed sites revealed that the fawns still
managed to find some cover in those habitats as well
(ARABLE: 4.8 6 7.8%, PASTU: 24.6 6 38.7%,
CLEAR: 34.5 6 40.5%). The best cover at the
random sites was found in YOUNG (23.4 6 47.8%)
and CON 5 (24.2 6 58%), but at the selected bed sites
the amount of canopy cover exceeded 24% in all
habitats, except ARABLE.
The relationship between visibility and canopy cover
We found a significant and negative relationship
between canopy cover and visibility at the random
bed sites (R2 ¼0.059, df¼76, P¼0.034) while no such
relationship appeared in the selected sites (R2 ¼
0.012, df ¼ 76, P ¼ 0.172). This means that high
canopy cover and low visibility covary at the habitat
level. However, fawns seem to select the two bed-site
variables (canopy cover and visibility) independently
from each other, but always better than random bed
sites.
Discussion
Neonate mortality of fallow deer was 23.6% and the
mortality caused by predation was low, since only
one of eight non-surviving fawns died from what we
defined as predation. As expected, fawns selected bed
sites that offered better concealment (lower visibility)
Table 2. Bed-site selection of fallow deer fans at Koberg in southwestern Sweden, during the summer of 2009. Based on a total number of 77
selected bed sites from 23 marked fawns ( þ 77 random bed sites). Surviving fawns: N¼18 and non-surviving fawns: N¼5. Relative (%) habitat
use is indicated within brackets. On arable land, three bed sites were situated in the edge zone (within 5 m) of forest. On clear cut, one bed site
was situated in the edge zone (within 5 m) of arable land. In forest, eight bed sites were situated in the edge zone (within 5 m) of arable land.
Environmental variables
Selection of bed sites
Total
number
ARABLE
PASTU
CLEAR
YOUNG
CON 5
CON 15
MIXED
WETLAND
25
5
6
5
5
19
12
0
Total
77
Habitat
Mean visibility (m)
Mean canopy cover (%)
%
Surviving
fawns
Non-surviving
fawns
Selected
bed sites
Random
bed sites
Selected
bed sites
Random
bed sites
32.5
6.5
7.8
6.5
6.5
24.7
15.6
0
20 (30.3)
4 (6.1)
6 (9.1)
5 (7.6)
2 (3.0)
18 (27.3)
11 (16.7)
0
5 (45.5)
1 (9.1)
0
0
3 (27.3)
1 (9.1)
1 (9.1)
0
3.1
1.7
1.3
2.4
8.2
6.9
4.3
NA
4.6
4.2
5.6
4.3
10.0
15.4
7.3
NA
4.8
24.6
34.5
47.8
58.0
29.8
30.2
NA
0.6
0.6
0.2
23.4
24.2
4.1
6.3
NA
66 (100)
11 (100)
100
286
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Figure 4. Mean visibility of selected bed sites compared to mean
visibility of random bed sites (6 SD) of each main habitat type for
fallow deer fawns in southwestern Sweden, during summer 2009.
Figure 5. Mean canopy cover of selected bed sites compared to
random bed sites (6 SD) of each main habitat type for fallow deer
fawns in southwestern Sweden, during summer 2009.
and more canopy cover than available at random
sites. As predation was low, we did not find support
for difference in bed-site selection between surviving
and non-surviving fawns. The high level of selectiveness when choosing a bed site, with no apparent effect
on fawn survival, does not entirely rule out neither of
the two bed-site selection hypotheses, i.e. the ’ghost
of predators past’ (Byers 1997a) or for thermoregulation (Linnell et al. 1995). We interpret our results to
be in line with Byers (1997a) hypothesis of ’pleiotropy’, i.e. an anti-predatory behaviour that remains
because it also has an alternative function (Stearns
2010). Being in an environment with few predators, it
seems as the anti-predator behaviour in our study
area is an adaption to a higher predation pressure
that have persisted under relaxed selection with
thermoregulation as the most likely alternative
function maintaining the behaviour.
Preliminary data indicating 30% neonate mortality in our study area is based on the difference in the
proportion of females with fetuses in the spring and
the proportion of lactating females next fall (P.
Kjellander, unpubl. data). This suggests that the
mortality rate of 23.6% in our study is not an
overestimation of the actual rates caused by capture
induced mortality. Six of the fawns died from
starvation. In all these cases, the necropsy showed
that none of these fawns had suckled for some time as
no signs of milk were found in the intestinal tracts.
We cannot exclude the possibility that the handling
and marking of the fawns had an effect on this, i.e.
causing the abandonment and subsequently starvation. However, studies of marking induced neonate
mortality in fallow deer (Galli et al. 2008) and other
ungulate species in enclosures (e.g. pronghorn
Antilocapra americana (Byers 1997b) and whitetailed deer (Ozoga & Clute 1988)) has shown no
evidence of increased mortality due to handling.
Furthermore, suckling usually occurs within the first
hour after birth (Chapman & Chapman 1997), and
since most of the fawns were estimated to be at least
24 hours or older at capture, they should have had
time to feed at least once before the possible
disturbance by the handling, suggesting that something was not right prior to the capture.
To our knowledge there are unfortunately no
previous studies on mortality rates in wild neonate
fallow deer. However, our estimated mortality rate of
, 24% is not particularly high in comparison to
studies of other species in predator-free environments but rather low for an area with predators
(Andersen & Linnell 1998). Gill (1994) found a mean
mortality rate of 26% for roe deer fawns, in a 23-year
data set from England in a predator-free area, and in
studies of other free-ranging ungulates in predatorfree environments, neonate mortality has been
reported to range between 10 and 20% (red deer
(Guinness et al. 1978), white-tailed deer (Ozoga
&Verme 1986, McGinnes & Downing 1977), moose
(Stubsjoen et al. 2000)). In contrast, a mortality rate
of . 40% is repeatedly reported for roe deer fawns in
areas were mainly red fox function as a predator of
fawns (Aanes & Andersen 1996). Our estimated
predation rate therefore seems very low in the light of
those reported on roe deer. It is likely that even
during the short period a fallow deer fawn is likely to
be killed by a red fox, the fox is too small a predator
to be as successful in killing fallow deer fawns of more
than double the size of a roe deer fawn, particularly as
it seems as fallow deer females actively and collec-
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287
tively successful defend their fawns. Roe deer
females, which are smaller than fallow deer females,
have been shown to be quite capable of defending
their fawns from red fox predation, either by
deterring the fox from areas were fawns are bedded
or by attacking and chasing the fox away (Jarnemo
2004). This suggests that female fallow deer would
have a good chance of defending their fawns as well.
In fact, repeated observations of females running
towards the field worker was done, and if the fawn
screamed during the handling, several females sometimes came running from different directions as an
obvious effect of the alarming fawn. This behaviour
indicates a strong willingness to defend fawns against
predators. In line with that, we once actually observed a group of females aggressively chase away a
fox from a fawn.
If the mortality rate reported in our study in large
is not an effect of handling or predation, why do the
fawns die? Veterinary autopsies, did not reveal any
acute diseases or high parasite loads. Even if no
extreme weather occurred during the two years, the
fawns did experience some heavy summer rains and
hypothermia can not be ruled out as the ultimate
death cause. Considering that the mean birth date of
fallow deer is 3-6 weeks later in our study area than
reported for the native deer species (moose (Sæther &
Heim 1993), red deer (Meisingset 2003) and roe deer
(Nordström et al. 2009)), this might indicate sensitiveness in fallow deer fawns to bad weather conditions.
On the other hand, studies of the relationship
between longevity and reproductive success in roe
deer (Kjellander et al. 2004) and reindeer (Weladji et
al. 2006) have demonstrated that females that live the
longest have the highest fitness. An age effect on
successful reproduction, attributed to the improved
maternal skills of older, more experienced females,
has also been recorded for pronghorn (Byers 1997a)
and white tailed deer (Ozoga & Verme 1986).
Furthermore, maternally inexperienced females are
suggested to be more likely to abandon their fawns
than older females (Ozoga & Verme 1986). Thus, a
reason for some of the recorded abandonments in
our study might be inexperience and possibly a
higher sensitivity to disturbance of young females.
The high population density in our study area could
further increase the losses in at least two different
ways. First, it is likely that this population is severely
food limited, and it is well established that females in
poor body condition give birth to fawns with low
birth weights which results in low survival (Guinness
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et al. 1978). Secondly, disturbance of human origin
and interactions of intra- or interspecific origin might
disturb the female to fawn imprinting process and
negatively affect the willingness of particularly young
inexperienced females to continue to invest in a fawn
also after a short separation, as observed in sheep
Ovis aries (Otal et al. 2009).
Albeit mortality biased towards males is often seen
in sexually dimorphic species (Owen-Smith 1993),
and more male fawns than female fawns actually died
during our study in both absolute (five males and
three females) and relative numbers (32% mortality
in males and 16% in females), the small sample size
could not with any significance confirm this as a
general pattern. To reach significant differences in
gender specific survival of that magnitude would
have required more than three times as many marked
fawns as marked in our study.
The spatial behaviour of fallow deer neonates was
examined at two different levels, i.e. the habitat
selection within their home range, which in fact is a
selection made by the mother, and the bed-site
selection within that habitat. Open habitats, including arable land and pastures, were selected over
forest habitats by fallow deer mothers at Koberg and
arable land was most used according to the CA. This
habitat also had the lowest canopy cover compared
to the other habitat types even though the measured
visibility to predators was quite low at these sites.
Since the habitat selection is made by the mother
within her home range, she has to consider her own
needs as well as the fawn’s (Ciuti et al. 2006), and one
of the most important factors generally affecting
habitat use is the availability and quality of forage
(Putman 1988). Being intermediate mixed feeders
(Hofmann 1989), fallow deer are known to spend
most of their time feeding in open grasslands
(Chapman & Chapman 1997, Borkowski & Pudelko
2007), and a previous study at Koberg (Winsa 2008)
confirmed that arable land was the preferred habitat
by fallow deer also in this study area. This supports
the findings in our study, as female fallow deer in our
study area did not make any trade-offs in their
habitat selection, but in fact used the habitat that
gained themselves in terms of energy intake, contrary
to fallow deer females in Italy that moved to poorer
but safer habitats when rearing offspring (Ciuti et al.
2006). However, even though canopy cover were low
in the selected habitat (arable land), fawns that
bedded in this habitat still seemed to have a good
concealment since the visibility was quite low at the
selected bed sites.
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Apart from the measured environmental variables, there could also be other factors explaining
habitat choice made by the mothers. Since fallow
deer females seem quite capable of defending their
fawns from red fox predation, they may also choose
to keep their fawns in open habitats in order to be
able to discover potential predators in time, suggesting that fallow deer females do make habitat choices
to reduce predation risk to their neonates. Furthermore, the female also needs to consider her own
safety and be able to detect other larger predators.
Since most of the fawns in our study were found
and marked in open habitats, this could be causing a
bias towards fawns using open habitats, giving the
possibility that the results of our study does not
reflect the true spatial distribution of the population.
However, even though habitats within the home
range were not used by random, all habitat types
were to some extent utilised by fallow deer mothers
and their neonates. This could imply that all habitats
were suitable for the fawns to find satisfying cover.
Additionally, as suggested by Linnell et al. (1999),
this could also be an anti-predator strategy by not
having a strong preference towards a single habitat
type, but instead utilising all habitat types, and
therefore forcing a potential predator to search all
available habitats, making the hiding strategy even
more effective.
Many bed sites of surviving fawns were located in
forest habitats with relatively high visibility. While
these habitats do not seem to provide protected bed
sites, the vegetation in the forest might still provide a
good concealment for the fawns when moving with
their mothers. This is in contrast with the open
habitats were the visibility of the fawns are low when
they are bedded, but when they are standing up and
moving to another location they might be spotted
from long distances. All habitat types except wet
lands were utilised for bed sites.
Despite predation pressure at our study site being
low, which was probably partly due to a high level of
predator control, the results of our study show that
there is still a risk for predation on fallow deer
neonates since one fawn most likely was killed by a
red fox. Furthermore, regardless of the fact that we
did not find any differences in the bed-site selection of
surviving compared to non-surviving fawns in our
study, the random bed sites could be thought of as
reflecting the habitat choice made by the female,
suggesting that females which choose a habitat with
low visibility gives her fawn a higher probability of
survival.
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Even though the low occurrence of predators
might explain the lack of importance of bed-site
visibility for the fawn survival, this does not
explain the fact that we did not find canopy cover
to be of any importance for the survival. In
accordance with the study of Linnell et al. (1999)
on roe deer fawns, we found that fallow deer fawns
selected bed sites that offered better canopy cover
than available at random sites. We did not,
however, find any evidence of the presumed
relationship between canopy cover and survival.
On the other hand, we found a significant and
negative relationship between canopy cover and
visibility at the random bed sites while no such
relationship appeared in the selected sites. We
interpret this as, while not surprisingly, high
canopy cover and low visibility covary at the
habitat level, fawns seem to select the two bed-site
variables (canopy cover and visibility) independently of each other. In a study from France, Van
Moorter et al. (2009) found that survival of young
roe deer fawns was positively related to the
selection of bed sites with more canopy cover,
but older fawns were rather selecting bed sites with
more light penetration, i.e. less canopy cover.
More importantly, they also report a connection to
weather conditions, where bed sites with denser
cover were used on days with low temperature and
bed sites with more light penetration on warm
days, thus fawns might search for exposure to sun
on sunny days (Van Moorter et al. 2009), perhaps
to dry up after rainfall or cold and humid nights to
reduce hypothermia. Thus, it makes us interpret
the role of canopy cover for fawn survival to be
partly in support of Byers (1997a) hypothesis of
’pleiotropy’. Following that, the significance of
thermoregulation will perhaps be the most important contemporary selection force in the absence of
large predators, since this behaviour could also
depend on weather conditions not investigated in
our study.
Acknowledgements - we are grateful to the Silfverschiöld
family for allowing us to work at their estate, to Anders
Friberg for his patience with all interruptions in his daily
work and to all field workers for making our study possible
by helping us to catch fawns. We are also grateful to
Guillaume Chapron for help with calculations in R and
Gustaf Samelius for assistance with ArcGIS and two
anonymous reviewers for comments and suggestions. Our
study was part of a larger project studying roe deer and
fallow deer interactions and the ecology of wild fallow deer in
Sweden managed by Petter Kjellander and was supported
by grants from the private foundation of ’Oscar och Lili
289
Lamms Minne’ to Ida Svartholm, The Swedish Environmental Protection Agency to Petter Kjellander and Ulrika
A. Bergvall, The Swedish Association for Hunting and
Wildlife Management to Petter Kjellander and Anders
Jarnemo.
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