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J. Zool., Lond. (2003) 261,119–128 C
2003 The Zoological Society of London Printed in the United Kingdom DOI:10.1017/S0952836903004059
Home range and habitat use by cheetahs (Acinonyx jubatus)
in the Kruger National Park
L. S. Broomhall1*, M. G. L. Mills1,2,3and J. T. du Toit1
1Mammal Research Institute, Department of Zoology & Entomology, University of Pretoria, Pretoria, 0002, South Africa
2South African National Park, Private Bag X402, Skukuza, 1350, South Africa
3Endangered Wildlife trust, Private Bag X11, Parkview, 2122, South Africa
(Accepted 6 March 2003)
Abstract
Cheetah Acinonyx jubatus home-range size and habitat use were analysed using radio-tracking data collected in
the southern district of the Kruger National Park (KNP) between 1987 and 1990. Meaningful estimates of home-
range size, using the 95% minimum convex polygon method, were 126 km2for a three-male cheetah coalition,
195 km2for a solitary male, and 150 km2and 171 km2for two female cheetahs. Although cheetahs used all
habitats according to their availability, they did show a preference for open savanna habitat because their core or
total home ranges centred on these habitats. Female cheetahs used denser woodland habitat more frequently than
males, as they seemed to be influenced by the distribution of their main prey, impala Aepyceros melampus,which
also preferred denser woodland habitat.
Key words:Acinonyx jubatus,cheetah, habitat, home range, impala, Kruger National Park
INTRODUCTION
Cheetahs Acinonyx jubatus are known as predators with
apreference for open plains habitats (Dorst & Dandelot,
1970). This is not only because they are built for speed
(Nowell & Jackson, 1996; Mills & Hes, 1997) but also
because most previous studies on cheetah ecology were
conducted in open grassland savannas, with a particular
focus on the Serengeti Plains in East Africa (Schaller,
1972; Frame & Frame, 1980; Caro & Collins, 1986,
1987; Durant et al., 1988; Fitzgibbon, 1990; Caro, 1994;
Laurenson, 1994, 1995a,b;Laurenson, Weilbnowlski &
Caro, 1995; Durant, 1998, 2000a,b;Kelly et al., 1998).
Cheetahs, however, also occur across a wide range of
woodland savannas (Myers, 1975; Skinner & Smithers,
1990; Mills & Hes, 1997), though in comparison far less
is known about cheetah ecology and behaviour in these
areas. This is largely because of the logistical constraints
associated with tracking and observing cheetahs in
wooded habitats.
Recent studies in woodland savannas have increased
our understanding of cheetah ecology in these areas (Zank,
1995; Hunter, 1998; Purchase & du Toit, 2000). To further
extend the documented information on cheetah ecology
∗All correspondence to: L. S. Broomhall School of Life & Environmental
Sciences, George Campbell Building, University of Natal, Durban 4041,
South Africa.
E-mail: l broomhall@hotmail.com
in woodland savannas we present a study conducted on the
home-range size and habitat use of cheetahs in the Kruger
National Park (KNP), South Africa. Cheetahs have never
been studied in the KNP before and being an important
conservation area for cheetahs, it will provide information
useful to park management. This will also assist with
cheetah conservation across a broader habitat scale as
woodland savannas make up a large part of the cheetah’s
range (Skinner & Smithers, 1990).
In this study, the habitat requirements of the cheetahs
in a woodland savanna are considered. In the Serengeti,
adequate concentrations of Thomson’s gazelle Gazella
thomsoni and sufficient cover were the main determinants
of male cheetah territory location as these features
attracted females (Caro, 1994). Cover is required for
stalking (Cohen, Scholtz & Reichel, 1978; Fitzgibbon,
1990; Caro, 1994), concealment from other predators and
resting (Caro, 1994; Zank, 1995; Purchase, 1998), and
would therefore be an important requirement for cheetahs
in this open grassland habitat. In contrast, in woodland
habitats, where there is considerable cover available, we
hypothesize that cheetahs would seek out open areas to
meet their hunting requirements. This is because they
require open spaces for high-speed chases as bushes and
trees may obstruct their hunting strategy (Myers, 1975;
Bertram, 1979; Broomhall, 2001). Therefore, assuming
that cheetahs need more open areas to hunt successfully,
we predict that in woodland savannas they will prefer open
habitats.
120 L. S. BROOMHALL,M.G.L.MILLS AND J. T. DU TOIT
Sabie and Crocodile rivers
Habitat types
Combretum woodland
Riverine thickets
Combretum / Terminalia woodland
Acacia thickets
Open savanna
Lebombo Hills
Kruger
National
Park
10 0 10 20
N
km
Fig. 1. Location of study areas in the Kruger National Park showing
six different habitat types.
METHODS
Study area
The field study was conducted in the southern district of
the KNP (24◦96–25
◦44E, 31◦30–32
◦00S) between the
Sabie and Crocodile rivers (Fig. 1). The southern district
covers an area of c. 3786 km2(Bowland, 1994). Two focal
study areas were located in this district: (1) the main
focal study area in the south-eastern region (6 radio-
collared cheetahs were tracked in this area); (2) a second-
ary focal study area to the west of the main study
site in a more central region of the southern district
(1 cheetah was tracked in this area). The KNP study area
lies in a summer rainfall region, with a mean annual rain-
fall averaging 600 mm rising to 700 mm in the Lebombo
Hills (Gertenbach, 1980).
The main study area comprises 3 broad habitat types,
identified using the landscape system developed by
Gertenbach (1983) (Fig. 1). The main landscape is classi-
fied as Sclerocarya birrea/Acacia nigrescens tree savanna
(an area covering c.250 km2)occurring on fairly flat undu-
lating terrain (Gertenbach, 1983). It is an open to semi-
wooded savanna with a moderate shrub layer and dense
grass layer which is intersected by several well-defined
and broad (50–200 m) drainage lines (Gertenbach, 1983;
Funston, 1999). The sides of the drainage lines are lined
with a denser shrub and tree layer than the rest of the open
savanna.
The Lebombo Hills border the open savanna to the east,
covering an area of c.148 km2.This is an undulating,
broken landscape with north–south rhyolite ridges and
bottomlands, 100 m higher than the basalt plains in
the open savanna (Gertenbach, 1983). The vegetation is
heterogeneous dense to moderate bush, dominated by
Combretum apiculatum,with a less dense field layer
(Gertenbach, 1983).
The landscape bordering the open savanna to the west
is the Acacia welwitschii thickets on Karoo sediments
(170 km2)described as dense thorny bush thickets
(Gertenbach, 1983). The structure of the woody com-
ponent is a moderate tree savanna with tall shrubs and
sparse low shrubs (Gertenbach, 1983). The grass cover
is less dense and sometimes disappears in the dry season
(Gertenbach, 1983).
The banks of Sabie and Crocodile rivers, which cut
through all 3 landscapes in the main study area, are densely
overgrown with woody species, and the grass layer is
usually absent (Gertenbach, 1983).
The secondary study area comprises a further 3 broad
habitat types (Fig. 1). The thickets of the Sabie and
Crocodile rivers (1148 km2)are low-lying, relatively flat
areas, characterized by dense woody vegetation, with
A. nigrescens/C. apiculatum dominating (Gertenbach,
1983). The Combretum collinum/C. zeyheri woodland
(454 km2)and mixed Combretum spp./Terminalia sericea
woodland (257 km2)are undulating landscapes on granite
with distinct uplands and bottomlands (Gertenbach,
1983). In both habitat types, the uplands have relatively
dense bush savanna, the bottomlands are open savanna
with a dense grass layer, while dense riverine vegetation
line the banks of drainage lines and rivers (Gertenbach,
1983).
Data collection
Seven adult cheetahs were radio-tracked for 3 years
between 1987 and 1990 (data collection by MGLM)
in the southern district of the KNP (see Table 1). All
cheetahs, except M3, were darted from a vehicle by slowly
approaching close enough to a distance of c.15–20 m.
M3 was trapped in a 2-door cage trap set at a latrine
with the access closed off by a thorn bush barrier around
the tree so that the only way the cheetah could get there
was through the trap. A rag dipped in urine from an
oestrus female in captivity was hung in the trap (Mills,
1996). Cheetahs were immobilized with 150–200 mg CI-
744 Park Davis (a 1:1 mixture of tiletamine hydrochloride
and zolazepam hydrochloride), or a combination of 75 mg
CI-744 and 15 mg xylazine and fitted with Telonics MOD-
315 collars weighing 100 g, except for M3 who was fitted
with a MOD-400 collar weighing 170 g. The duration of
the tracking period for each cheetah varied depending
on circumstances. M1 was only tracked until his collar
stopped transmitting, M2 was found dead after both he and
his companion had contracted sarcoptic mange, and the
collar from M3 was removed. F1’s collar was removed, F2
could not be relocated after her radio stopped transmitting
after just over 1 year, F3 disappeared and is believed to
have died, as did F4, who might have emigrated as she
wasayoung adult when caught.
14697998, 2003, 2, Downloaded from https://zslpublications.onlinelibrary.wiley.com/doi/10.1017/S0952836903004059 by The University Of Newcastle, Wiley Online Library on [05/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Home range and habitay use by cheetahs 121
Tab l e 1 . Home-range estimates (km2)ofradio-tracked cheetahs Acinonyx jubatus in the southern district of the Kruger National Park
using the minimum convex polygon method (MCP). M3, three-male cheetah coalition; M2, two-male cheetah coalition; M1, single male
cheetah; F1–F4, female cheetahs
Estimates of home-range size
No. of Asymptote
Cheetahs Period tracked fixes reachedaMCP 100% MCP 95% MCP 50%
M3 25 Feb 87–15 Apr 90 177 Yes 170 126 14.1
M2 11 Jul 88–14 May 89 21 No 438 242 b
M1 14 Oct 88–02 May 90 27 Yes 261 195 21.5
F1 29 Nov 88–17 Apr 90 70 Yes 192 150 36
F2 16 Sep 87–07 Oct 88 25 Yes 179 171 35.7
F3 18 Aug 87–23 Oct 87 14 No 118 118 b
F4 29 Aug 89–28 Apr 90 9 No 102 102 b
aNumber of fixes was plotted against home-range size to determine if home-range size reached an asymptote (Harris et al., 1990;
Kenward & Hodder, 1996).
bSample sizes too small to estimate 50% MCP.
Three types of data collection were used based on
the duration of the observation period: (1) radio-location
observations, when only a radio-fix of the animal was
recorded; (2) short-term continuous observations, when
radio-collared cheetahs were followed by vehicle for
periods of 2–15 h; (3) long-term continuous observations,
when cheetahs were followed continuously for 14 days. All
3data collection types were recorded for a 3-male cheetah
coalition (M3) and a female cheetah (F1). Two 14-day
observation periods were recorded for M3 and 1 14-day
period was recorded for F1. Radio-locations only were
recorded for a 2-male cheetah coalition (M2); a solitary,
territorial male cheetah (M1); and 3 female cheetahs F2,
F3 and F4. M2 and F4 were dispersing, sub-adult cheetahs.
All cheetahs used all or part of the main study area,
except cheetah M1, who inhabited the central region of
the southern district of the KNP. The dispersing, sub-adult
male cheetah coalition (M2) and sub-adult female cheetah
(F4) were radio-collared in the main study area, but ranged
westwards into the central region. During the entire study,
only 1 other cheetah, an adult male, observed once, was
seen in the main study area. In 1987, M3, F2 and F3 the
main study area, and in 1988 and 1989 it was inhabited by
M3, F1, F2 and F4.
Home range
The home-range sizes of 7 radio-collared cheetahs were
determined using an ArcView extension package Animal
Movement (Hooge, 1999). Location points from radio-
tracking and direct observations recorded between 1987
and 1990 were used for home-range analyses. The location
points were recorded on an old grid reference system used
by the KNP and converted to latitude and longitude for
input into the models. Models were run using the Universal
Transverse Mercator (UTM) co-ordinate system. Only
location points taken c.24h apart were used to ensure
independence of locations (Swihart & Slade, 1985). The
number of fixes needed to calculate home-range size
were analysed using the computer package RangesV
(Kenward & Hodder, 1996) by plotting number of fixes
against home-range size until home-range size reached an
asymptote (Harris et al., 1990).
Two non-parametric methods were chosen to estimate
home-range size. The minimum convex polygon (MCP)
method (Jenrich & Turner, 1969) is the most widely
used method in the literature (Harris et al., 1990) and is
presented for comparison with other studies. The biggest
problem with this technique is that area and shape are
heavily influenced by outlying fixes (Harris et al., 1990)
and may include large unused areas. To address these
problems, the peeled minimum convex polygon method
was used to remove ‘outliers’ or fixes showing ‘excursive
activity’ (Mizutani & Jewell, 1998), because Burt (1943)
considered excursions outside the normal area not part of
an individual’s ‘normal’ home range. Animal Movement’s
harmonic mean method for outlier removal was used to
plot the 50% and 95% MCPs (Hooge, 1999). The 50%
and 95% home ranges were selected as they are generally
considered the most robust estimators of an animal’s centre
of activity (core home range) and a close approximation
of total range size, respectively (Jaremovic & Croft, 1987;
Harris et al., 1990; White & Garrott, 1990; Mizutani &
Jewell, 1998).
Habitat use
Achi-square goodness of fit test (Zar, 1999) was used
to determine if the observed frequencies of habitat use
differed significantly from expected frequencies based
on the proportion of area contributed by each habitat
within a cheetahs’ home range. Radio-locations recorded
during direct observations of 4 radio-collared cheetahs
(M3, M1, F1 and F2) were used to determine frequencies
of observed sightings per habitat type. Distinct vegetation
types defined at the landscape level (according to
Gertenbach’s landscapes, see KNP study area) are referred
to in this paper as habitat types. The 95% MCP was
used to delineate home ranges for calculating habitat
availability (km2)for individual cheetahs, as this method
was considered to provide the best estimates of home range
for these cheetahs. Habitat analyses were not conducted
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122 L. S. BROOMHALL,M.G.L.MILLS AND J. T. DU TOIT
on female cheetahs F3 and F4 owing to small sample sizes
and dispersing male cheetahs (M2).
Impala were the cheetahs’ main prey in the study
area, and their habitat use was also analysed using the
same methods. The mean number of impala per hab-
itat type was determined using KNP aerial census data
collected every August between 1987 and 1990 (Joubert,
1983). Habitat availability for impala in the main study
area was calculated by measuring the total area of each
habitat type lying between the Sabie and Crocodile
rivers. Bonferroni confidence intervals were performed
thereafter, for those use/availability comparisons that were
found to be significantly different in the chi-squared tests,
to determine preference or avoidance of particular habitat
types (Neu, Byers & Peek, 1974; Byers & Steinhorst,
1984) by cheetahs and impalas.
Differences in habitat use by the 3-male coalition (M3)
and female cheetahs (F1 and F2) were investigated using
chi-square test on contingency tables (Zar, 1999). Further
differences were investigated within habitat types, as the
sides of the drainage lines intersecting these habitats had
thicker shrub and tree cover: 2-sample t-tests were used to
test for differences in mean distance from all male and
female cheetah locations to the nearest drainage line
and road.
Impala herd locations, obtained from aerial census data
collected during dry and wet months between 1986 and
1990 within the main study area, were digitized into
ArcView. Using ArcView and a grid overlaying the study
site (grid scale c.4km
2per cell), the frequencies of
cheetah and impala herd locations within cheetah home
ranges (using the 100% MCP as this included all cheetah
fixes) were counted per grid cell. Frequencies of impala
herd sightings per grid cell were averaged for wet and
dry months. Spearman rank correlation (Zar, 1999) was
then used to test for relationships between distributions of
impala herds and cheetahs (M3 and F1) in wet and dry
seasons.
Whilst recording activities during direct observations
of cheetahs M3 and F1, relative grass height and shrub
cover were also recorded: grass height was classified as
(1) short <20 cm, (2) medium 20–60 cm, (3) tall >60 cm;
shrub cover was classified as: (1) open, (2) moderate,
(3) dense (Funston, Mills & Biggs, 2001). Nine different
cover classes were created by all possible combinations of
grass height and shrub cover, e.g. short grass, moderate
bush, etc. A chi-squared test on a contingency table was
used to determine if the cheetahs showed preferences for
any vegetation classes based on the vegetation observed
at the start of each observation period. To satisfy sample
size requirements per cell for the chi-square test (Roscoe
&Byars, 1971), the vegetation classes were further
combined into 4 categories of relative cover ranging from
habitats with open to closed cover: (1) short grass, open to
moderate shrub cover; (2) medium grass, open to moderate
shrub cover; (3) short to medium grass, dense shrub cover;
(4) tall grass with any shrub cover.
The amount of time a cheetah spent walking, resting
and hunting within the different shrub cover classes were
also recorded for the same 14-day observation periods.
Although the availability of each vegetation class could
not be quantified, this was used as indication of habitat
use by male and female cheetahs (observations could not
be tested statistically owing to small sample sizes).
The frequencies of scent markings by the 3-male co-
alition (M3) along the roads and off the roads were
recorded during a 14-day continuous observation period.
The relative importance of roads to the coalition was ana-
lysed using a chi-square goodness-of-fit test by comparing
the observed frequency of scent markings along and off
the roads to the expected frequency based on the total area
of all roads (averaging 6 m wide) vs ‘off the roads’ area
available within the home range of the cheetah coalition.
RESULTS
Home range
An asymptotic home range was not reached for female
cheetahs with <25 fixes (Table 1). These were also
considerably smaller than the other female cheetah
home ranges and were therefore not considered accurate
estimates (Table 1). The two-male cheetah coalition (M2)
ranged widely and only 21 fixes were obtained for these
animals, which may explain why an asymptote was not
reached for their home range (Table 1). Additionally, these
animals were young and had not acquired a territory.
Cheetah home ranges using the 100% MCP method
are presented in Figs 2 & 3 as this method could be
used to represent all cheetah home ranges regardless of
sample size or male social status. With the exception of
the dispersing sub-adults F4 and M2, the home ranges
of cheetahs centred on the open S. birrea/A. nigrescens
tree savanna in the main study area (Fig. 2), which also
made up a large percentage of their home ranges (Table 2).
The territorial three-male coalition (M3) had smaller total
and core home-range sizes than adult female cheetahs with
adequate sample sizes (Table 1). There was a large amount
of overlap between adult cheetah home ranges in the main
study area (Fig. 2), although there was no overlap in core
home ranges between M3 and F1 and some overlap with
F2 (Fig. 4). The core home ranges of all the cheetahs were
located in the open savanna, although the core ranges of
both female cheetahs bordered onto and extended into the
Lebombo Hills (Fig. 4). The single male cheetah (M1)
radio-tracked in the central region of the southern district
(Fig. 3) had a far larger home range than the three-male
cheetah coalition in the open savanna (Table 1). Most of
the home range of M1 was positioned in the Combretum
and Combretum/Terminalia woodlands (Fig. 3). The sub-
adult two-male coalition was first located in the open
savanna of main study area, but dispersed westwards,
moving over greater distances than all the other cheetahs
(Fig. 3, Table 1).
Habitat use
Chi-square analyses showed that the observed habitat use
by female cheetah F2, the single male cheetah M1 and
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Home range and habitay use by cheetahs 123
Female cheetahs
Male cheetah coalition
Rivers
505km
N
A
BC
F2
F1
F4
F3
Lower SabieLower SabieLower Sabie
Fig. 2. Home range of female cheetahs Acinonyx jubatus (F1, F2,
F3 and F4) and a three-male cheetah coalition M3 in the south-
eastern region of Kruger National Park, estimated by the 100%
minimum convex polygon (MCP) method. A, Acacia thickets;
B, open savanna; C, Lebombo Hills.
Tab l e 2 . Percentage of habitat within the home range of a cheetah
Acinonyx jubatus in the south-eastern region of the Kruger National
Park using the 95% minimum convex polygon (MCP) method
%incheetah’s home range
Habitat M3 F1F2F3
Acacia thickets 10 15 8 10
Open savanna 86 69 63 90
Lebombo Hills 4 16 29 –
male cheetah coalition M3 did not differ significantly
from that expected based on habitat availability within
their home ranges (95% MCP). The female cheetah F1,
however, showed a significant difference between the
observed and expected habitat use (χ2=9.6, d.f. =2,
P<0.05) (Table 3). The Acacia thickets were used
more than expected and the open savanna and Lebombo
Hills less than expected. Bonferroni confidence intervals,
however, indicated that F1 showed no significant
differences between observed verses expected utilization
of available habitat types. The expected value for the
Acacia thickets (Table 3) may be the cause of the
discrepancy as the Chi-squared goodness of fit test can
be sensitive to small values (Zar, 1999). The observed
habitat use by impala differed significantly from the
Two-male cheetah coalition
Single male cheetah
Rivers
50 5 km
N
Lower SabieLower SabieLower Sabie
10
E
F
D
E
D
AB
C
Fig. 3. Home range of a single male cheetah Acinonyx jubatus M1
and two-male cheetah coalition M3 in the southern district of the
Kruger National Park, estimated by the 100% minimum convex
polygon (MCP) method. A, Acacia thickets; B, open savanna;
C, Lebombo Hills; D, riverine thickets; E, Combretum/Terminalia
woodland; F, Combretum woodland.
Tab l e 3 . Chi-squared test for use of different habitat types by the
cheetah Acinonyx jubatus female F1 in the south-eastern region of
the Kruger National Park
Habitat Observed Expected
Acacia thickets 10 4
Open savanna 46 50
Lebombo Hills 11 13
expected, based on habitat availability between the Sabie
and Crocodile rivers (χ2=185, d.f. =2, P<0.0001).
Bonferroni confidence intervals indicated that the impalas
preferred the Lebombo Hills, avoided the open savanna,
and used the Acacia thickets in proportion to its
availability.
Habitat use by male and female cheetahs, based on the
frequency of locations per habitat type, was significantly
different (χ2=25.75, d.f. =2, P<0.0001). While the
number of observed locations of the male cheetah
coalition (M3) was greater than the expected in the open
savanna and less than expected in the Acacia thickets
and Lebombo Hills, the number of female cheetah (F1)
locations was greater than expected in the Lebombo Hills
and Acacia thickets and less than expected in the open
savanna. The number of female cheetah (F2) locations
was less than expected in the open savanna and greater
than expected in the Lebombo Hills.
When plotting radio-locations in the main study area,
the distribution of fixes revealed that the three-male
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124 L. S. BROOMHALL,M.G.L.MILLS AND J. T. DU TOIT
Male cheetah coalition M3
Female cheetah F1
Sabie River
km202
Female cheetah F2
N
B
A
C
Fig. 4. Core home ranges of male (M3) and female cheetahs
Acinonyx jubatus (F1 and F2) in the south-eastern region of the
Kruger National Park, estimated by the 50% minimum convex
polygon (MCP) method. A, Acacia thickets; B, open savanna;
C, Lebombo Hills.
Roads
N
Drainage lines Acacia thickets Open savanna Lebombo Hills
Fig. 5. Radio-location points of a three-male cheetah Acinonyx jubatus coalition M3 (䊉)andfemale cheetahs F1 and F2 (䉱)inthe
south-eastern region of the Kruger National Park.
cheetah coalition M3 was closely associated with the
roads, while female cheetahs (F1 and F2) followed the
drainage lines (Fig. 5). Unpaired t-tests showed M3
was significantly closer to the roads than the drainage
lines (t=2.42, d.f. =352, P<0.05), while F1 and F2
were significantly closer to the drainage lines than
the roads (t=4.64, d.f. =138, P<0.0001 and t=2.55,
d.f. =48, P<0.05, respectively; Fig. 6). The mean
distance to drainage lines and roads between males
and females were also significantly different (Fig. 6).
Female cheetahs F1 and F2 were significantly closer to
the drainage lines than the male cheetah coalition M3
(t=3.92, d.f. =245, P<0.0001 and t=4.37, d.f. =200,
P<0.0001, respectively), and M3 was significantly closer
to the roads than F1 (t=3.6, d.f. =245, P<0.001), but
not F2.
There was a significant positive correlation between
impala herd and female cheetah F1 locations in both the
wet (rs=0.401, P<0.05, n=39) and dry (rs=0.488,
P<0.01, n=39) seasons. No correlations were found
between impala herd and male cheetah (M3) locations
at any time (wet: rs=0.161, n=43; dry: rs=−0.189,
n=43). There was a strong significant difference between
the frequency of scent markings by M3 along and off the
roads, based on the area available for scent marking within
their home range (χ2=13736, d.f. =1, P<0.0001). The
frequency of scent markings was greater than expected
along the roads (observed 265, expected 5) and less than
expected off the roads (observed 52, expected 312).
Chi-squared analysis based on frequency of observa-
tions in different vegetation-cover categories showed the
three-male cheetah coalition to be using significantly
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Home range and habitay use by cheetahs 125
M3 M3 F1 F1 F2
0
250
500
750
Mean distance (m)
To roads To drainage lines
Cheetahs
P< 0.05
P< 0.001
P< 0.05
F2
Fig. 6. Mean distance ( ±SE)ofmaleandfemale cheetah Acinonyx
jubatus locations to drainage lines and roads in the south-eastern
region of the Kruger National Park. M3, three-male cheetah
coalition; F1 and F2, female cheetahs.
different vegetation categories for different activities
(χ2=26.1, d.f. =6, P<0.01). The coalition preferred
medium to tall grass areas for hunting, but not in areas
with dense shrub cover. Short to medium grass regardless
of shrub cover was preferred for resting, while moderate
to more dense shrub cover was preferred for walking,
but not areas with tall grass. Female cheetah F1 did not
show any preferences for different vegetation categories
for different activities.
The percentage of time recorded in different shrub
cover classes indicated some differences between the male
cheetah coalition M3 and female cheetah F1. The female
spent 72% of her time in moderate shrub, 14% in dense
shrub and 14% in open shrub, while the males spent 51%
of their time in moderate shrub and 49% in open shrub.
DISCUSSION
Estimates of home-range size for cheetahs in the KNP
were presented in this study (Table 1). Bowland’s (1994)
photographic survey estimated that the home-range size
of male and female cheetahs in the KNP varies between
104 km2and 1848 km2.These estimates, however, were
taken from small sample sizes and the social status of
the animals was unknown. Therefore, non-territorial or
dispersing cheetahs may be partly responsible for the
large home ranges recorded in Bowland’s (1994) study.
When compared with other studies in other woodland
savannas, cheetah home-range sizes varied widely from
24 km2in Matusadona National Park (MNP) to 320 km2
in Kgalagadi Transfrontier Park (KTP) (Mills, 1998;
Purchase & du Toit, 2000; Broomhall, 2001). The
variation in cheetah home-range size, including those
observed between M1 and M3 in this study, may also
be associated with the variation in habitat structure and
prey availability across different landscapes or habitat
types. The resource dispersion hypothesis predicts that
territory size is determined by the dispersion pattern of
food patches (Macdonald, 1983) or other resources. In
the Serengeti, the large female home ranges (averaging
833 km2)are attributed to the patchiness of their main
prey, not low prey density, as Thomson’s gazelle undertake
large seasonal migrations (Caro, 1994). Similarly, the
nomadic nature of springbok Antidorcas marsupialis in
the KTP (Mills, 1998) and concentrated distribution of
high prey density on the foreshore grassland in MNP
(Purchase & du Toit, 2000) may have influenced the
home-range size of female cheetahs in these areas.
Similarly, the distribution of suitable habitat for hunting
may affect cheetah home-range size. Kruuk (1986) states
the distribution of such places would be more important
in determining the size of home ranges in felids, than
the number of potential prey moving around an area. In
Phinda Resource Reserve (PRR), cheetahs sought out
open grassland patches in the woodland for hunting,
which constituted only 8.6% of available habitat and
were distributed as small, discreet, widely spaced patches
(Hunter, 1998). Hunter (1998) considered that suitable
habitat for hunting, rather than prey, may have been the
‘patchy’ resource that gave rise to cheetah movements.
In this study, total and core territory sizes of the three-
male cheetah coalition M3 were smaller than the adult
female home ranges (Figs 2 & 4, Table 1). Although it
is unusual in carnivores for the territory size of males to
be smaller than the range size of females (Sandell, 1989;
Mizutani & Jewell, 1998), increasing female home-range
size is associated with reduced male territory size, since
very large female areas are no longer defensible by males
(Caro, 1994). This is true in the Serengeti (Caro, 1994) and
is probably the situation in the KNP as female cheetahs
have large home ranges. Cheetah home ranges in the main
study area of this study overlapped extensively (Fig. 2) and
the territory of the male cheetah coalition overlapped those
of both adult female cheetahs, presumably to increase
opportunities for mating (Sandell, 1989; Caro, 1994).
However, core home ranges of the male coalition M3 and
female F1 did not overlap and may not have overlapped
with female F2 had the sample size for F2 been >25
(Table 1, Fig. 4). Thus, although home ranges showed
extensive overlap, the cheetahs also seemed to avoid each
other and showed exclusive use of certain areas, which
is more typical of asocial felids (Caro, 1994; Mizutani &
Jewell, 1998).
Although chi-squared analyses showed that cheetahs
had no preferences for any particular habitat types, the
home ranges of adult cheetahs in the main study area
were centred on the open savanna (Fig. 2, Table 2),
suggesting a preference for this relative to other available
habitats. The core home ranges of cheetahs M3, F1 and
F2 were also positioned in the open savanna (Fig. 4).
Pienaar (1969) recorded that cheetahs showed a decided
preference for open or lightly wooded savannas across
the KNP. In the Timbavati and Klaserie private nature
reserves, where the habitat varies from open savanna
to moderately dense or riparian woodland, cheetahs also
preferred the A. nigrescens/S. birrea woodland (Kruger,
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126 L. S. BROOMHALL,M.G.L.MILLS AND J. T. DU TOIT
1988). The solitary male cheetah in the central study
area, positioned his territory in the Combretum woodlands
(Fig. 3), which is characterized by open savanna habitat in
the bottomlands (see Methods). In the woodland savanna
of PRR, the core areas of the territorial male cheetahs
with the longest tenure were all centred in regions with
the most extensive grasslands and all females used these
same grasslands extensively (Hunter, 1998). In MNP,
home ranges of cheetahs included both woodland and
grassland habitat, where they preferred the open grassland
for hunting (Purchase & du Toit, 2000).
When comparing the habitat selection of male and
female cheetahs, the females showed greater use and pre-
ference than males throughout for denser vegetation types,
such as the drainage lines and more woody Lebombo
Hills and Acacia thickets. Impala also preferred these
dense vegetation types (Gertenbach, 1983; Ben-Shahar,
1995; this study). Therefore, females may be using thicker
vegetation because encounters with impala are greater.
This is supported by the significant positive correlation
found between distribution of female cheetah F1 and the
distribution of impala, while no correlations were found
between impala and the male coalition M3. In the KNP,
the open savanna is an important habitat to the cheetahs as
it is the preferred habitat for hunting (Broomhall, 2001).
Therefore, it seems that female cheetahs must reach a
compromise between the habitat best suited for hunting
and the one supporting the highest impala densities.
Wooded areas are nevertheless preferred for walking
and moving between hunting and resting sites (Cohen
et al., 1978; Zank, 1995; Purchase, 1998). The three-
male cheetah coalition selected moderate to dense shrub
cover for walking and female F1 seemed to use denser
shrub cover for all activities. This may be because the
cheetahs are avoiding predators by selecting habitats that
provide greater concealment. In the Serengeti, Durant
(1998) found that cheetahs seek out ‘competitive refuges’
with low densities of lions Panthera leo and spotted hyenas
Crocuta crocuta,asthey are both directly responsible for
cub mortalities (Laurenson, 1994) and both species steal
kills from cheetahs (Caro, 1994). The risk of cub predation
may be another reason why the core ranges of female
cheetahs bordered onto and extended into the Lebombo
Hills (Fig. 4) because female cheetahs were accompanied
by cubs in over two-thirds of observations. This habitat not
only carries higher impala densities (Broomhall, 2001),
butisalso avoided by lion and spotted hyena (Mills &
Biggs, 1993; Mills & Gorman, 1997). The Acacia thickets,
on the other hand, are preferred by lion and spotted hyena
(Mills & Biggs, 1993) and carry high densities of these
predators (Mills & Gorman, 1997).
Unlike the female cheetahs in the KNP, the male
coalition was less concerned with cover and impala
distribution. They seemed to prefer the more open savanna
habitat, used more open vegetation when resting compared
to other activities, and concentrated their activities along
the roads. The male coalition may be less concerned
than the females about other predators as a cheetah
coalition is more likely to deter other predators than a
solitary animal (Caro, 1994). Male coalitions also prey
on larger food items (Caro, 1994; Broomhall, 2001) and
seem less dependent on impala than female cheetahs
in the KNP (Broomhall, 2001). In addition, staying in
open habitat might make it easier for them to detect
intruding cheetah males. The greater dependency of
female cheetahs on impala may be because of the high
nutritional demands placed on females during pregnancy,
lactation and cub growth (Laurenson, 1995b). Kruuk
(1986) suggests that females must exploit a less profitable
butmore predictable food supply, i.e. impala, than the
males, which may indicate that females in the KNP are
distributing themselves according to impala distribution
and are not only avoiding predators.
On the Serengeti Plains, where the males position their
territories and where there is some cover, Caro (1994)
considered that the availability of sufficient cover for
stalking and resting determines territory location. In this
study, we consider that scent-marking may also influence
the habitat selection of territorial male cheetahs. Scent
marking plays an integral role in territory maintenance as
awarning to other males of their presence (Eaton, 1970),
butisalso considered important to males and females for
communicating their reproductive status (Marker-Kraus
et al., 1996). Scent marks are usually placed on cons-
picuous objects or frequently used places where encounter
rates are maximized (Gorman & Trowbridge, 1989), such
as large trees or shrubs and dirt mounds (Eaton, 1970), and
roads or game paths or around waterholes (Funston, 1999).
The male cheetah coalition in the KNP scent marked
significantly more frequently along the roads and MGLM
observed that they preferred the large trees on the sides of
roads rather than the bushes for scent marking. Cheetahs
on the open grassland plains (Caro, 1994) use prominent
landmarks throughout their territory, such as solitary trees,
rocks and termite mounds far from others (Caro, 1994).
In Nairobi National Park, Eaton (1970) found that the
distance between scent marking locations halved with
greater densities of woody plants, indicating the need for
trees or bushes for scent marking. Cheetah scent marking
behaviour and the importance of habitat for scent marking
and territorial advertisement, however, remains relatively
unexplored.
Our study in the KNP found that cheetahs prefer more
open habitat in woodland savannas. However, for female
cheetahs this preference seems to be largely influenced
by the distribution of impala. Predation may also play
asignificant role in habitat selection but this could not
be determined in this study. The need for open areas
for hunting may be the main determinant for cheetahs
using the open savanna, which indicates the importance
of open habitats to cheetahs. In Uganda, a GIS analysis
of vegetation structure in areas where cheetahs were
observed and in those where none were reported suggested
that cheetahs favoured habitats with 25–50% woody cover
(Gros & Rejm´
anek, 1999). The requirements by cheetahs
for open areas in woodland savannas is important as the
effects of dense or encroaching bush may lower cheetah
densities and affect population viability in small reserves.
Finally, we concede that due to difficulties associated
with tracking and observing cheetahs in this study the
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Home range and habitay use by cheetahs 127
cheetah observations were limited to small sample sizes.
This study, however, provides a meaningful contribution to
our knowledge of cheetah ecology in woodland savannas,
which to date has been largely unexplored. Further
studies in woodland savanna ecosystems are required to
substantiate any differences in habitat use by male and
female cheetahs and to gain a better understanding of the
effects of prey availability and their requirements for open
areas and cover on home-range size and habitat use.
Acknowledgements
The National Research Foundation, Endangered Wildlife
Trust, University of Pretoria and Mammal Research
Institute generously provided funding for this research.
Many thanks to the late Ross English and to Bryan
Havemann, Heather Wildi and Adriaan Louw for assisting
with field work. Thanks to Human Buirski for help on
all IT problems; Judith Kruger and other KNP staff for
responding to many GIS requests; Martin Srauss for
advice on RangesV; Ian Meiklejohn and Gaby van Wyk
for help with GIS ArcView; Jaco Barendse for proof-
reading the manuscript on several occasions; and Pat
Fletcher, Martin Haupt and Elmarie Cronje for assisting
with numerous logistical and administrative concerns.
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