ChapterPDF Available

An adaptive management approach to trophy hunting of leopards (Panthera pardus): a case study from KwaZulu-Natal, South Africa


Abstract and Figures

Leopards have considerable value as a trophy hunting species in South Africa and the national CITES quota was recently doubled from 75 to 150 leopard hunts per year. This decision was taken despite a lack of reliable data on regional leopard population abundance and with little understanding of the possible impacts of increased harvesting on the species. In KwaZulu-Natal (KZN), we showed that an uneven distribution of hunting effort contributed towards high mortality rates and low recruitment in a nominally protected leopard population. Against this backdrop we developed a revised protocol for the trophy hunting of leopards in KZN based on five key recommendations: (1) limiting the number of CITES tags allocated in KZN to five each year, (2) ensuring a more even distribution of permits across the province, (3) allocating applications for leopard hunts to individual properties rather than hunting outfitters, (4) linking the likelihood of obtaining a tag to the size of the property, and (5) restricting the trophy hunting of leopards to adult males. This protocol and, as such, has potential to act as a model for the hunting of the species where currently permitted or proposed across its range.
Content may be subject to copyright.
An adaptive management approach
to trophy hunting of leopards
(Panthera pardus): a case study
from KwaZulu-Natal, South Africa
Guy A. Balme, Luke T.B. Hunter, Pete Goodman, Hayden Ferguson,
John Craigie, and Rob Slotow
A radio-collared female
leopard in the Phinda-
Mkhuze ecosystem, South
Africa. #Brett Pearson.
McDonald 14-McDonal-chap14 Revises Proof page 341 22.10.2009 10:52am
In principle, the trophy hunting of large carnivores
has substantial potential to foster their conservation
(Lindsey et al. 2007; Loveridge et al. 2007b). Predators
may be tolerated to a greater extent when they aug-
ment human livelihoods, and hunting may be a
means by which private landowners and local com-
munities can generate revenue from the presence of
carnivores (Leader-Williams and Hutton 2005). If the
profits realized from harvesting a few individuals are
sufficient incentive for people to tolerate the larger
population, the goals of trophy hunting and conser-
vation are compatible. However, poorly managed tro-
phy hunting may produce a variety of deleterious
effects on carnivore populations. Excessive and sus-
tained harvesting of a species can lead to its local
extinction, or reduce numbers to such an extent that
a population is no longer viable in the long term
(Treves and Karanth 2003). Similarly, even when quo-
tas for trophies appear conservative, hunting may con-
stitute additive rather than compensatory mortality,
particularly given that other anthropogenic sources of
mortality are rarely included in the calculation of quo-
tas (Caro et al. 1998). Additionally, uncontrolled tro-
phy hunting may erode the genetic diversity of a
species by consistently selecting the fittest individuals
in a population that would normally survive to propa-
gate the species, particularly adult breeding males and
females (Harris et al. 2002; Coltman et al. 2003). Final-
ly, hunting can alter demographic patterns in carni-
vore populations resulting in impacts on non-hunted
cohorts (Tuyttens and Macdonald 2000). Accordingly,
for trophy hunting to be considered a legitimate tool
in the management and conservation of carnivores, it
is essential that hunting practices avoid detrimental
impacts on populations and are demonstrated to be
biologically sustainable.
The African leopard is one of the most sought-after
big game trophies (Turnbull-Kemp 1967; Grobbelaar
and Masulani 2003). Although resilient in the face of
human pressure, leopards have been eradicated from
vast tracts of their former range due mainly to loss of
habitat, depletion of natural prey, and direct perse-
cution by people (Nowell and Jackson 1996). Leo-
pards once occurred throughout most of the
African continent with the exception of the hyper-
arid interiors of the Sahara and Namib deserts (Fig.
14.1). They are now virtually extinct in North Africa,
have disappeared from most of the West African
coastal belt, and continue to decline outside of pro-
tected areas in large parts of East and southern Africa
(Hunter et al., in press). Ray et al. (2005) estimate that
leopards have been eradicated from 36.7% of their
historic African range. Notwithstanding these recent
declines, leopards are often deemed to warrant low
conservation priority. Their wide geographic range
and ability to persist in regions where other large
carnivores have been extirpated has given rise to a
widespread assumption that their conservation
status is assured. Although listed on appendix I of
the Convention for the International Trade in
Endangered Species (CITES) of flora and fauna, 12
African countries are permitted to export a quota of
leopard skins procured through trophy hunting
(CITES 2007). In 2008, the quota for all states com-
bined was 2648 leopard skins/year, with an addition-
al 10 skins from the Democratic Republic of Congo
and Gabon obtained from sources other than hunt-
ing (Table 14.1).
Trophy hunting is regulated and accounts for a
smaller proportion of leopard deaths than other
human-mediated mortality such as problem animal
control and illegal persecution; for example, an aver-
age of 445 leopards were legally hunted per year in
the 1980s compared to an estimated minimum of
770 killed by other anthropogenic sources in South
Figure 14.1 Current (dark grey) and historical leopard
distribution. Modified from Rayet al. 2005, with permission.
342 Biology and Conservation of Wild Felids
McDonald 14-McDonal-chap14 Revises Proof page 342 22.10.2009 10:52am
Africa, Botswana, Zimbabwe, and Zambia combined
(Martin and de Meulenaer 1988). Nevertheless, there
is remarkably little scientific input on the allocation
of harvest quotas and the implementation of hunt-
ing practices. Population estimates most widely
quoted by countries proposing to increase or intro-
duce quotas are derived from an overly simplistic
modelling exercise that correlated leopard numbers
with rainfall (Martin and de Meulenaer 1988). This
model was widely criticized for omitting critical
factors such as anthropogenic mortality and prey
availability, and for relying upon questionable as-
sumptions, for example, that leopards occur at maxi-
mum densities in all available habitats. Accordingly,
the final estimate of 714,000 leopards in sub-Saharan
Africa was considered an impossible overestimate
(Jackson 1989; Norton 1990; Bailey 2005). Despite
recent advances in survey techniques that furnish
accurate estimates of leopard numbers at moderate
cost (Henschel 2001; Balme et al., 2009a), few autho-
rities employ these techniques in routine manage-
ment activities such as setting hunting quotas.
Despite the lack of data, the demand for leopard
hunting in Africa appears to be growing. The annual
quotas of four countries (Namibia, South Africa, Tan-
zania, and Mozambique) have increased since 2002,
and Uganda recently introduced trophy hunting of
the species, raising the combined quotas of these
countries from 485 to 1048 (CITES 2007). Further
potential for over-exploitation exists where regula-
tions are abused or ignored. In Tanzania, females
comprised 28.6% of 77 trophies shot between 1995
and 1998, even though only males are legally hunted
there (Spong et al. 2000a). In Zimbabwe, excess skins
from illegal hunts are reportedly smuggled into Zam-
bia and Mozambique to be exported under the quo-
tas of those countries (I. A. Caldwell, personal
communication). As an explicit condition of CITES
certification, African countries that allow leopard
hunting commit to do so in a way that is compatible
with national and sub-national conservation objec-
tives for the species. However, there remains (1) no
rigorous data on the numbers, densities, or popula-
tion trends of leopards anywhere they are hunted;
(2) few empirical data on the impact of hunting on
leopard populations; and, finally, (3) no national,
provincial, or local regulatory framework for harvest-
ing leopards established by an assessment and con-
sideration of numbers 1 and 2.
A variety of management strategies have been pro-
posed to regulate trophy hunting of comparable car-
nivore species that have similar lacunae in
knowledge (Logan and Sweanor 1998; Laundre
Clark 2003). In lieu of detailed demographic data,
any sensible approach must incorporate an under-
standing of the basic population dynamics of the
target species. In particular, the source–sink meta-
population structure of large felid populations has
been used as a proxy to partition areas into sub-
populations, each with distinct management objec-
tives (Cougar Management Guidelines Working
Group 2005). In this context, source areas are closed
to hunting and function as inviolate refuges where
natural population dynamics can occur not directly
affected by human influence. They serve as robust
biological savings accounts that ensure long-term
population persistence and provide dispersing ani-
mals for numeric and genetic augmentation of
neighbouring sinks and other, more distant sources
(Weaver et al. 1996). Sink areas, in contrast, are those
sub-populations where harvesting is permitted. They
are generally, but not always, less suitable for carni-
vores due mainly to the presence of people, and
concomitant ecological changes such as reduced
Table 14.1 The annual CITES quotas for the 12
African countries permitted to export leopard skins
procured through trophy hunting in 2008.
Country Quota
Botswana 130
Central African Republic 40
Ethiopia 500
Kenya 80
Malawi 50
Mozambique 120
Namibia 250
South Africa 150
Tanzania 500
Zambia 300
Zimbabwe 500
Uganda 28
Trophy hunting of leopards 343
McDonald 14-McDonal-chap14 Revises Proof page 343 22.10.2009 10:52am
prey availability and habitat quality (Cougar Man-
agement Guidelines Working Group 2005).
We adopted a similar approach to develop a novel
protocol for the trophy hunting of leopards in Kwa-
Zulu-Natal (KZN), South Africa. We previously de-
monstrated that poorly regulated harvesting
contributed to the low reproductive output of leo-
pards in the Phinda-Mkhuze ecosystem, one of the
largest leopard populations in the province (Balme
and Hunter 2004; Balme et al., 2009b). In that con-
text, we developed a new protocol for the trophy
hunting of leopards in KZN, intended to be compati-
ble with both the conservation objectives for the
species as defined by the statutory conservation au-
thority, Ezemvelo KwaZulu-Natal Wildlife (EKZNW),
and the desire of private landowners and the hunt-
ing industry to utilize leopards. The chief objectives
of the new hunting protocol were to (1) mitigate the
detrimental effects of leopard hunting in KZN, (2)
enhance both the sustainability and economic pro-
ductivity of leopard hunting in KZN, and (3) ensure
compliance with international and national norms
and standards for leopard hunting (Department of
Environmental Affairs and Tourism, South Africa
2006). In this chapter, we review the historical back-
ground of leopard hunting in KZN, summarize the
biological impacts of hunting on a local leopard
population, and present the new protocol for trophy
hunting the species in the province.
Trophy hunting of leopards in
KZN province is situated on the eastern side of South
Africa (26!450–31!100S and 28!450–32!500E;
Fig. 14.1) in the biologically rich transition zone be-
tween the Indian Ocean and the Drakensberg escarp-
ment. Although it is only the sixth largest (92,100
) of South Africa’s nine provinces, it contains the
most people (estimated at 9.5 million in August 2001;
Statistics South Africa 2005). Altitude ranges from sea
level in the east to over 3450 m above sea level in the
west. Vegetation types vary from tropical dune forests
near the coast to moist lowland and upland grass-
lands, a variety of subtropical forests, and semi-arid
savannas further inland, all of which contain the
megafauna typical of these habitats in Africa, though
many now occur only in protected areas (Goodman
2003). Approximately 58% of the land area in KZN is
used for stock farming (including of wild game), 17%
for crops, 8% for commercial forestry, and roughly 7%
is set aside for conservation (South African National
Biodiversity Institute 2005).
The most suitable leopard habitat in KZN is found in
the mesic northeastern parts of the province on com-
mercial game ranches, cattle farms, and within state-
managed protected areas. Leopardsalso occur on com-
munally owned Zulu lands but generally at lower den-
sities chiefly due to habitat degradation and lack of
natural prey (Balme and Hunter 2004). We refer to
both private and communal land as ‘properties’. Le-
gally, leopards are categorized as ‘specially protected
game’ and fall under the mandate of EKZNWon pub-
lic, private, and communal lands. They are strictly
protected inside provincial public parks (IUCN Pro-
tected Areas Management Category II), and all trophy
hunting of the species currently takes place on private-
ly owned land. Otherwise, leopards cannot legally be
destroyed, unless they represent a threat to life or
property, and then only if the affected party has ap-
plied for and been granted a ‘destruction permit’ by
EKZNW. Despite these restrictions, leopards are killed
opportunistically and illegally on private and commu-
nal lands in the province by livestock owners and
game ranchers (Balme et al., 2009b).
From 1996 onwards, the annual quota of CITES
permits allocated for trophy hunting in KZN has
varied between 5 and 10 leopards (Burgener et al.
2005). Since 2003, it has been stable at 5 animals/
year. By comparison, in 2006, Limpopo Province was
allocated 50 permits, North West Province received
20 permits, and the adjoining Mpumalanga Province
received 10 permits (Burgener et al. 2005). Prior to
2006, CITES permits were allocated in KZN using a
random draw process whereby the first five appli-
cants drawn received a permit and the remaining
applicants were placed sequentially on a waiting
list. If a leopard was not successfully hunted within
2 weeks, the permit defaulted to the next person on
the waiting list, and so on.
For such a widespread species, removing five ani-
mals from the provincial population does not seem
excessive. However, hunting effort was not evenly
distributed throughout KZN. Only 5 of the 22
344 Biology and Conservation of Wild Felids
McDonald 14-McDonal-chap14 Revises Proof page 344 22.10.2009 10:52am
conservation districts that fall within potential leop-
ard range in the province were awarded CITES per-
mits between 2000 and 2005 (Table 14.2). The
distribution of tags across these five districts was
also uneven, with most (79%) allocated to the neigh-
bouring Nyalazi and Mkhuze districts (Table 14.2),
and specifically to private land surrounding the
Phinda-Mkhuze Game Reserves, which constitute a
single contiguous leopard population. During the
same period, Nyalazi had more than double the
amount of leopards shot on CITES hunts than any
other district (Table 14.2). Sixteen properties in the
province were allocated CITES permits between 2000
and 2005; however, only seven successfully hunted
leopards. The number of leopards shot per property
during this period ranged from 1 to 4 (mean ¼2.29 #
1.21). Additionally, and contrary to national stipula-
tions (Department of Environmental Affairs and
Tourism, South Africa 2006), leopards were frequent-
ly hunted on the same property in consecutive years.
Three privately owned game farms hunted leopards
for 2 successive years and another game farm hunted
leopards for 4 consecutive years. None of these prop-
erties was larger than 25 km
, which is slightly less
than the mean home-range size of female leopards
resident in protected areas in the region (mean ¼
27.57 #5.72 km
; G.A. Balme, University of
KwaZulu-Natal, unpublished data).
This high offtake contributed to a significant local
effect on the leopard population. The average annual
mortality rate (AMR) of this population between
2002 and 2005 was 0.401 #0.070 (Balme et al.,
2009b), more than double that recorded for the spe-
cies in similar habitat under protection (Bailey 2005).
Male leopards suffered particularly high mortality
(AMR ¼0.538 #0.023), due partly to their higher
vulnerability to trophy hunting (Balme et al., 2009b).
Males are more desirable to trophy hunters because
of their larger size, and males also utilize larger home
ranges and cover greater daily distances than females
(Mizutani and Jewel 1998; Bailey 2005), increasing
their chances of moving into areas where they can be
hunted. In some felid and ursid species, unnaturally
high turnover among males can result in elevated
levels of infanticide (Swenson et al. 1997; Wielgus
and Bunnell 2000; Logan and Sweanor 2001; Whit-
man et al. 2004). Although infanticide is a natural
event in leopards (Ilany 1986; Scott and Scott 2003),
high levels of male turnover promotes a situation in
which females cannot successfully raise cubs because
of constant incursions by new males. The conse-
quences of this in the Phinda-Mkhuze population
included low survival rates among cubs (AMR ¼
0.574 #0.089), delayed age at first parturition
(45.33 #1.76 months), reduced conception rates
(19%), and low annual litter production (0.643 #
0.127 litters/female/year; Balme et al., 2009b). The
combined effects of high mortality and depressed
reproduction led to a negative population growth
rate during the study period (l¼0.978). It is impor-
tant to note that human hunting was not the only
source of leopard mortality and many losses in the
Table 14.2 The number of CITES permits allocated to, and the relative success of, hunts within conservation
districts in KwaZulu-Natal (KZN) province, South Africa, from 2000 to 2005.
Number of permits allocated Successful hunts
Conservation district N%n% Hunt success per district (%)
Dundee 1 2 0— 0
Mdletshe Tribal Authority 1 2 0— 0
Mkhuze 15 32 4 25 27
Nyalazi 22 47 9
56 41
Vryheid 8 17 3 19 38
Total 47 16 Mean = 34
In addition to this total, an adult female leopard was critically wounded in a hunt in the Nyalazi district in 2003 but not recovered; she is
unlikely to have survived.
Trophy hunting of leopards 345
McDonald 14-McDonal-chap14 Revises Proof page 345 22.10.2009 10:52am
population were the result of illegal persecution rath-
er than legal trophy hunting (Balme et al., 2009b).
Nevertheless, the concentration of trophy hunting
effort on a single population contributed to higher
AMRs than previously documented for the species,
prompting EKZNW to ratify the new protocol to-
wards the end of 2005.
Our protocol was based on five key recommenda-
tions intended to achieve objectives 1–3 listed earlier.
These recommendations, and the rationale behind
them, were as follows:
1. Cap the number of CITES permits allocated in
KZN to five each year. When South Africa’s an-
nual CITES quota of 75 leopards was doubled in
October 2004, 10 permits were made available
for trophy hunting in KZN. A population and
habitat viability analysis (PHVA) which evalu-
ated the possible effects of increasing the leop-
ard harvest suggested that the KZN population
had a much greater likelihood of suffering sig-
nificant impacts from hunting than did more
robust populations elsewhere in South Africa
(Daly et al. 2005). This was primarily because
the level of illegal killing of leopards by humans
was estimated among the highest in the coun-
try. According to the PHVA, the risk of extinc-
tion rose from 11% to 62% and the mean
population size declined from an estimated
393 animals to 217 with a quota of 10 leopards
(Fig. 14.2; Daly et al. 2005). Thus, despite the
relatively large estimated population size and
carrying capacity, the removal of just five addi-
tional leopards per year put this population at
substantially greater risk (Daly et al. 2005).
2. Allocate applications for leopard hunts to indi-
vidual properties rather than hunting outfitters.
In the past, any number of hunting outfitters
could apply to hunt leopards on the same prop-
erty, with potential for consecutive hunts dur-
ing the year and multiple animals being
removed from a single farm in a year. Under
the new protocol, landowners applied for a
hunt to take place on a particular property
(this applies whether the land is privately or
communally owned). If successful, the proper-
ty, not the property owner or hunting outfitter,
was allocated the leopard hunt. It then fell to
the property owner to negotiate the best deal
from an outfitter and inform EKZNW of the
name of outfitter and timing of the proposed
hunt so that the CITES permit could be issued.
3. Ensure a more even distribution of CITES per-
mits across the province. As discussed earlier,
CITES permits were formerly allocated uneven-
ly across the province where it was both feasible
to hunt leopards and where landowners sought
to host hunts. To distribute hunting opportu-
nities more evenly, we demarcated five leopard-
hunting zones (LHZ), each with a population
considered extensive and robust enough to sus-
tain hunting (Fig. 14.3). Each LHZ covered an
area of at least 600 km
and lay adjacent to a
protected area of at least 290 km
in which
hunting was prohibited. Based on density esti-
mates from mark–recapture statistics applied to
camera-trapping we conducted in the main
land use types in which leopards occur in KZN
(11.11 leopards/100 km
in protected areas,
7.17 leopards/100 km
in game ranches, and
2.49 leopards/100 km
in non-protected areas;
Balme et al., in review), we estimated that each
LHZ contained at least 25 adults and was adja-
cent to a protected source population with at
least 32 adults. Each LHZ qualified for a single
permit (i.e. only one animal can be hunted each
year) and had its own individual draw and wait-
ing list. A CITES permit allocated to one LHZ
could not be used in another LHZ, regardless of
demand, or of the success of hunts. If a leopard
was not shot on the first allocated property
within 1 month (rather than the previously
stipulated 2 weeks), the permit was moved to
the next property on the waiting list of that
particular LHZ, and so on.
4. Link the likelihood ofobtaining a tag to the size of
the property on which the hunt will take place.
Properties within an LHZ were still drawn at ran-
dom; however, the chance of a property being
selected was weighted according to its size. This
was based on the assumption that, in the same
area, larger properties are likely to sustain more
346 Biology and Conservation of Wild Felids
McDonald 14-McDonal-chap14 Revises Proof page 346 22.10.2009 10:52am
leopards. Therefore, for every 1.0 km
of land per
property, that property qualified for a single ‘tick-
et’ in the draw for that LHZ. Thus, a 5 km
ty was eligible for 5 tickets, a 25 km
property was
eligible for 25 tickets, and so on. In any one draw,
the probability of any property being selected
increased with its size. Once a property had been
allocated a CITES tag, all of its tickets were
removed before the next draw which established
the first property on the waitinglist, and so on.
5. Restrict the trophy hunting of leopards in KZN
to adult males. South Africa is one of the few
countries that have historically allowed the
hunting of female leopards. The PHVA demon-
strated that harvesting only males resulted in
lower extinction rates for all provinces but was
most beneficial in KZN, where the risk of extinc-
tion decreased by almost half (Daly et al. 2005).
Therefore, it was proposed that only adult male
leopards over 3 years old should be legally
hunted in KZN. At this age, male leopards are
easily distinguishable from females; the majori-
ty of 3-year-old males weigh over 60 kg (mean ¼
64.50 #3.7 kg and n¼10; G.A. Balme and L.T.
B. Hunter, Panthera, unpublished data) which
is considerably heavier than females (mean ¼
35 #3.5 kg and n¼7) and they show numerous
obvious physical differences (Fig. 14.4). All
hunters taking leopards in South Africa are re-
quired to be accompanied by a professional
hunter (PH) who is responsible for reporting
the hunt to EKZNW. An EKZNW officer is re-
quired to inspect the trophy within 24 h of the
hunt and, provided all legal stipulations are
met, issue a CITES tag so that the trophy can
be exported. It is the responsibility of the PH
conducting the hunt to judge whether the ani-
mal to be shot is an adult male. A printed guide
is provided by EKZNW (Balme and Hunter, in
press) to PHs and hunting clients to assist them
in sexing and aging animals so that the likeli-
hood of shooting females is reduced.
This protocol represents the first rigorous effort to
ensure the trophy hunting of leopards in South
Africa is sustainable. Prior to this, and presently else-
where in South Africa, CITES permits were allocated
on a relatively ad hoc basis that resulted in a concen-
tration of hunting effort on a single leopard popula-
tion. Our protocol addressed this problem by
distributing hunts over a much larger area and is
based on the meta-population management ap-
proach used to manage puma (Puma concolor) and
grizzly bear (Ursus arctos horribilis) populations in
North America (Servheen et al. 1999; Laundre
Clark 2003). All LHZs in KZN adjoin large protected
areas that potentially act as source populations to
replace animals shot on surrounding farms and
game ranches. The implicit assumption is that each
LHZ is capable of sustaining the removal of one leop-
ard a year to trophy hunting without threatening the
demographic or genetic viability of that sub-popula-
tion or the larger regional population. In other parts
of South Africa, source populations may need to be
identified that do not comprise formally protected
land. In this context, there is considerable scope for
the private sector to contribute; for example, by the
Annual CITES quota
PE (100 years)
90 (6) 120 (8) 150 (10)
135 (9)105 (7)75 (5)
Waterberg Free State
KZN E Cape Mountains
E Cape Valleys Wild Coast
Figure 14.2 Effects of CITES quota on probability of
extinction of leopard populations in South Africa (from Daly
et al., 2005, used with permission). KwaZulu-Natal quota in
Trophy hunting of leopards 347
McDonald 14-McDonal-chap14 Revises Proof page 347 22.10.2009 10:52am
consolidation of commercial game ranches and pri-
vate reserves that do not rely on consumptive utili-
zation of leopards into ‘conservancies’, in which
multiple, private landowners develop agreements to
manage and conserve their combined areas as single
unit (the ‘conservancy model’; Hunter et al. 2003).
However, it is critical that source areas are large
enough that losses in adjacent sinks do not have
detrimental effects on the source population. Deter-
mining the size of a source area depends on various
factors such as prey density, habitat quality, and har-
vest rates in neighbouring sink populations. Beier
(1993) and Logan and Sweanor (2001) suggested
that a minimum size of 1000–2200 km
was needed
to sustain a viable puma population for 100 years. In
KZN, only two protected areas (Hluhluwe-Imfolozi
Game Reserve and the Mkhuze/Mun-Ya-Wana/Ma-
khasa/St Lucia Complex) adjoining a LHZ were this
size, but leopards reached higher densities in these
areas than pumas in the above examples and all sub-
populations were likely large enough to withstand
the removal of a single animal by hunting.
Although sink areas in our protocol comprised
non-protected farm land, commercial game ranches,
and communal lands, leopard population growth
rate need not be negative and, if the management
objective is to maintain hunting, harvest levels must
not exceed immigration rates (Cougar Management
Guidelines Working Group 2005). Indeed, provided
other anthropogenic sources of leopard mortality are
curtailed, populations occupying game-ranching
areas have the potential for positive growth given
the widespread availability of natural prey and habi-
tat (Lindsey et al. 2005). This may benefit ranch
EKZNW protected areas
South Africa
Commerical game ranches
Leopard hunting zones
KZN provincial boundary
Figure 14.3 The location of
designated leopard-hunting zones
(LHZ) in relation to commercial game
ranches and Ezemvelo KwaZulu-Natal
Wildlife (EKZNW) protected areas and
conservation districts in KwaZulu-Natal
(KZN) Province, South Africa.
348 Biology and Conservation of Wild Felids
McDonald 14-McDonal-chap14 Revises Proof page 348 22.10.2009 10:52am
owners seeking to profit from hunting leopards but
will only be realized when removal of leopards for
other reasons (primarily by illegal shooting and
trapping in retribution for real and perceived losses
of livestock) are reduced. Most importantly, we do
not accept that a sink is an area where uncontrolled
removal of leopards is permissible. Private land-
owners who hunt leopards and benefit financially
from their presence must foster populations on
their lands if both hunting and leopards are to persist
in those areas. This may be assisted by our recom-
mendation of linking leopard hunting to the size of
properties. As well as rewarding larger landowners
with an increased opportunity of hosting a hunt,
this may encourage smaller landowners with an in-
terest in hunting leopards to increase the size of their
land or consolidate neighbouring properties, and
therefore foster the growth of larger contiguous
areas of suitable habitat for leopards.
While we believe we have been conservative in
planning LHZs, we have not addressed a prevailing
problem anywhere that leopard quotas are assigned,
which is the quantification of additional leopard
deaths due to people. Caro et al. (1998) warned that
high levels of traditional hunting by resident com-
munities and illegal poaching probably resulted in
unsustainable trophy hunting quotas for leopards in
Tanzania, though that country subsequently dou-
bled its quota to 500 in 2002. In KZN and South
Africa generally, leopards are illegally killed primarily
by pastoralists and farmers, though there is some
evidence for organized poaching; for example, skins
of at least 58 individuals were seized in the Mkhuze
district in July 2004, apparently destined for interna-
tional markets (Hunter et al., in press). During a
6-year period, 52.6% of all anthropogenic leopard
mortality (n¼19) recorded in the Nyalazi district
was illegal (Balme et al., 2009b). This is undoubtedly
a minimum estimate given that most illegal killings
are concealed and even accidental deaths of leopards
such as road deaths are rarely reported. The ongoing
sustainability of leopard hunting in each LHZ will
depend on increased quantification and mitigation
of the numbers of individuals killed illegally.
In addition to illegal persecution, significant num-
bers of leopards are legally destroyed every year
through problem animal control usually undertaken
by the landowner. In principle, landowners wishing
to destroy a damage-causing leopard must demon-
strate that a particular individual is preying upon
livestock or creating some other form of damage
and that steps have been taken to bring about a
non-lethal solution to the problem (Ferguson
2006). In practice, destruction permits are regularly
awarded on the basis of little evidence and there is no
mechanism to ensure that landowners remove the
individual responsible for the damage (G.A. Balme
and L.T.B. Hunter, unpublished data). Although re-
cent policy changes by EKZNW attempt to address
this (Ferguson 2006), the number of leopards
killed as problem animals (3.42 #1.21; EKZNW, un-
published data) generally exceeds the numbers
hunted legally every year and often occurs on
the same properties as trophy hunting. Ideally,
Figure 14.4 Extract from a guide for PHs to assist in the
accurate identification of suitable leopards for hunting
(G.A. Balme and L.T.B. Hunter, Panthera, unpublished
report): a) adult female leopard highlighting narrower
chest area and proportionally smaller head, and b) adult
( > 3-years-old) male leopard showing developed neck
region with prominent dewlap and much larger head.
Trophy hunting of leopards 349
McDonald 14-McDonal-chap14 Revises Proof page 349 22.10.2009 10:52am
damage-causing leopards could be hunted under the
CITES quota (Daly et al. 2005) but this is not a viable
option in KZN. Trophy hunts are planned months
and sometimes years in advance while stock-killing
incidents are largely unpredictable. With only five
hunts taking place during the course of a year, it is
unlikely that the two events will occur concurrently.
Additionally, for this to work, the allocation of CITES
permits would have to be linked to complaints rather
than by the existing lottery process. Such a system
might foster incentives for false claims about dam-
age-causing leopards to increase the chances of re-
ceiving a CITES tag. Similarly, landowners with a
genuine problem of losing livestock would be re-
warded, potentially promoting lax husbandry. It is
possible that this idea may be more applicable else-
where such as in the Limpopo Province where there
is both a larger quota and greater conflict between
leopards and farmers (Daly et al. 2005). In Namibia, a
‘Hunters’ Hotline’ was established to link farmers
that were losing livestock to leopards with the Pro-
fessional Hunters Association (Stein 2008). While
this partially overcame the problem of sourcing cli-
ents available at short notice, it did not encourage
farmers to improve their management techniques.
Furthermore, a questionnaire survey revealed that
only 12% of farmers were willing to use the hotline
(Stein 2008). Most farmers thought it was easier and
more beneficial to destroy the problem leopard
themselves. In Namibia, a destruction permit can
be obtained after a damage-causing animal has
been killed.
Our protocol was not created to disadvantage hun-
ters or prevent the consumptive utilization of leo-
pards. In fact, the new system may improve the
overall economic productivity of hunting as more
professional hunters and landowners stand to benefit
from the limited CITES permits available. Previously,
the industry was monopolized by a small number of
outfitters operating in the Nyalazi and Mkhuze dis-
tricts. By distributing permits evenly across the prov-
ince, it gives a larger number of outfitters the
opportunity of hosting a leopard hunt. Additionally,
extending the validity of a permit from 2 weeks to 1
month greatly improves the chance of any individual
hunt being successful without increasing the harvest
quota. Individual outfitters therefore stand to gain as
their clients are typically charged a daily rate and there
is now potential for longer visits. Similarly, there is
increased potential for more property owners to
benefit financially over time from having leopards
on their land. The prospect of receiving revenue from
a trophy hunt every few years may mitigate damage
caused by leopards and encourage landowners to
allow the continued presence of leopards on their
farms where formerly they stood no chance to host a
hunt. Thirty-nine percent of landowners interviewed
in the Nyalazi district (n¼18) supported the new
protocol, 11% felt that the allocation of CITES permits
should be linked to the control of damage-causing
leopards, 11% preferred the old process, and 39%
showed no interest in hunting leopards (Balmeet al.,
2009b). More widely, if applied correctly, this frame-
work may ultimately permit increased quotas and
hunting in more areas of the province. If landowners
benefiting from hunts reduce the persecution of leo-
pards and foster population growth on their lands, the
potential exists to increase quotas and expand or add
LHZs to new areas.
Under our protocol, the potential also exists for a
wider cross-section of the community to benefit from
leopard hunts. Historically, leopard hunting has been
restricted largely to private farms and game ranches
and has rarely taken place on communally owned
Zulu land. Hunting outfitters had little incentive to
develop wider relationships as they were guaranteed
access to a small number of private landowners mostly
within the Nyalazi and Mkhuze districts. Under the
new system, outfitters who negotiate with local tribal
authorities (TAs) to conduct hunts will increase their
chances of securing access because communal lands
are generally much larger than individual private
holdings and therefore stand a greater likelihood of
being drawn in the lottery. At least two TAs, the
Mdletshe TA and the Mthembeni TA, already have a
system in place for such negotiations and received a
CITES permit for a leopard hunt in 2001 and 2007,
respectively. Other communities have developed
structures for hunting other trophy species that
could easily be extended to include leopards. For ex-
ample, the Makhasa TA hosted a black rhino (Diceros
bicornis) hunt in 2006 that generated almost US
$11,000 for the community (EKZNW, unpublished
data). Communal areas in KZN comprise approxi-
mately 30,000 km
, much of it with considerable po-
tential to be important leopard habitat (McIntoshet al.
350 Biology and Conservation of Wild Felids
McDonald 14-McDonal-chap14 Revises Proof page 350 22.10.2009 10:52am
1996). This will only likely be realized when commu-
nities are encouraged to change current land-use prac-
tices that limit the viability of the land to harbour
leopards and other wildlife. With its low operating
costs and high profit margin, trophy hunting may
provide one such incentive (Lewis and Jackson 2005).
The implementation of our protocol is only the
start of an adaptive process that will rely upon careful
monitoring of leopard numbers and losses to people.
This currently takes place in only one LHZ as part of a
long-term ecological study on the species (Hunter
et al. 2003; Balme and Hunter 2004; Balme et al.
2009a, Balme 2009b). Our data from this study
suggest the new protocol has been successful in facil-
itating the recovery of the Phinda-Mkhuze leopard
population (Balme et al., 2009b). Since its implemen-
tation in 2006, estimated population growth has
increased by 16% (l¼1.136), annual mortality has
dropped to 0.134 #0.016, and the reproductive
output of the population has improved (Balmeet al.,
2009b). Specifically, mean age at first parturition has
decreased (33.67 #1.85 months), conception rates
have improved (39%), and annual litter production
has increased (0.709 #0.135 litters/female/year;
Balme et al., 2009b). Most notably, all cubs (n¼14)
born to radio-collared females survived to indepen-
dence after the change in hunting protocol. As a result
of lowered harvest rates and increased social stability,
population density increased from 7.17 #1.12
leopards per 100 km
in 2005 to 11.21#2.11 leopards
per 100 km
in 2009 (Balme et al., 2009b).
Although it is unlikely such intensive monitoring
could be extended to other LHZs, the process may be
assisted by recent improvements in survey meth-
odologies for cryptic species. Camera-trapping is
one such method that furnishes accurate density
estimates for leopards with modest financial re-
sources and relatively little technical expertise
(Henschel and Ray 2003; Balme et al., 2009a). The
monitoring of trophy quality and composition will
also help gauge whether there are any undesirable
demographic shifts within a population (Anderson
and Lindzey 2005). The new reporting requirement
for trophy age and sex will improve monitoring of
trophy quality across all LHZs in KZN. In Botswana
and the Niassa Province in Mozambique, assess-
ments of trophy quality are used to promote sustain-
able leopard hunting (Begg and Begg 2007; Funston,
personal communication). Quotas are assigned inde-
pendently to regions depending on the sex and age
of animals taken in the previous hunting season.
These systems are self-regulating and encourage eco-
logically sound hunting by penalizing hunters that
shoot females or under-age leopards. Although no
similar penalties are currently in place for hunters
in KZN and the hunting of males only (recommen-
dation 5) is simply advised as best practice, more
stringent control is likely in the near future; for ex-
ample, a policy is being drafted that will prevent PHs
receiving a CITES permit for a period of 3 years if they
hunt female or subadult (<3 years old) leopards
(EKZNW, unpublished data). Evaluating and incor-
porating the socio-economic attitudes of human
communities coexisting with leopards will also be
crucial in successfully implementing this protocol.
Aside from anecdotal information (e.g. Hunter et al.
2003), there are no data available on the attitudes of
local communities towards leopards and their man-
agement, nor on the underlying factors influencing
attitudes. Practical solutions exist for conducting
widespread surveys in the region (Lindsey et al.
2005) and should be adopted more widely in parallel
with the implementation of the new protocol in all
Our protocol has potential to act as a standard for
hunting the species in other provinces of South
Africa, and further afield in Africa. Currently, no
other national or regional authority assigns quotas
based on a detailed understanding of the species’
ecology and the potential impacts of hunting as we
have attempted here. Implementing this approach
on a national scale will require that local conserva-
tion authorities clarify leopard population status and
distribution within each province of South Africa. As
a first step, we recommend the delineation of leopard
sub-populations that could act as source or sink areas
utilizing GIS coverages that include habitat and land-
use type, records of leopard presence, patterns of
problem-leopard complaints, historical utilization
patterns of leopards, and the demand for CITES tags
(Cougar Management Guidelines Working Group
2005). Although parameters will vary, this approach
is relevant to most countries that permit sport hunt-
ing of leopards. Similarly, our approach may have
value for other large, wide-ranging felids that are
sport-hunted, for example, cheetahs (Acinonyx
Trophy hunting of leopards 351
McDonald 14-McDonal-chap14 Revises Proof page 351 22.10.2009 10:52am
jubatus) in Namibia which are hunted exclusively on
private land (CITES 1992), pumas in Latin American
populations where sport hunting is growing (Ojasti
1997), and Eurasian lynx (Lynx lynx) in Norway
where quotas are assigned on the basis of depreda-
tion complaints (Andre
`net al. 2006). Except for a
handful of sites where lions have been reintroduced
(Hunter et al. 2007c), the leopard is the apex predator
in many landscapes in KZN and, as such, fulfils a
vital role in functioning ecosystems. We believe the
implementation of this protocol will be a key factor
in maintaining leopards in those landscapes by bal-
ancing the ecological requirements of the species
with the needs of multiple stakeholders.
We are grateful to all at EKZNW who helped to de-
velop and implement this protocol, and to the vari-
ous professional hunters, game-ranch managers,
farmers, and landowners who have agreed to adopt
it in the field. We are thankful to CC Africa/&Beyond
and the Wetland Authority for allowing us to con-
duct the research on leopards in their reserves. We
are especially grateful to Kevin Pretorius (CC Africa/
&Beyond) for inviting us to initiate research into the
effects of hunting on leopards in this region, and for
continued logistic and material support. This study
was supported by Albert and Didy Hartog, Panthera,
Christian Sperka and the Wildlife Conservation Soci-
ety. G.A. Balme was supported while writing this
chapter by a Panthera Kaplan Award and received a
NRF bursary from grant #FA2004050400038. Nicole
Williams is thanked for her considerable effort
in proofreading and formatting the manuscript.
We thank David Macdonald, Michael J. Chamber-
lain, Cynthia Jacobson, and five anonymous re-
viewers whose detailed comments improved the
chapter. Our thanks to Brett Pearson for providing
the photograph of Ntombi and to S. Nijhawan
(Panthera) who prepared the maps.
352 Biology and Conservation of Wild Felids
McDonald 14-McDonal-chap14 Revises Proof page 352 22.10.2009 10:52am
... A key underpinning for efficient and reliable population monitoring of large felids and other mammalian species is the ability of researchers to identify individuals that make up a wild population (Kuningas et al., 2014;Balme et al., 2019). This is crucial because individual misidentifications may adversely affect management decisions, conservation assessment and even lead to erroneous quota setting for extractive activities like trophy hunting (Balme et al., 2010). Since the ground-breaking work of Karanth (1995) and Karanth and Nichols (1998), survey methods that aid in individual identification are commonly followed by the capture-recapture (CR) framework to estimate their populations. ...
... This is because practitioners are required to devote more time and effort to reliably identify individuals which lack distinct markings (Mattioli et al., 2018) or by choosing a survey method that can detect individuals based on other features. Failure to adopt appropriate survey methods increases the chances of individual misidentification (Soller et al., 2020), may inflate or deflate capture rates Garrote et al., 2021), bias sex/age-class structure of the population (Balme et al., 2012), and adversely impact population management and conservation decisions (Balme et al., 2010). ...
Full-text available
Large felids represent some of the most threatened large mammals on Earth, critical for both tourism economies and ecosystem function. Most populations are in a state of decline, and their monitoring and enumeration is therefore critical for conservation. This typically rests on the accurate identification of individuals within their populations. We review the most common and current survey methods used in individual identification studies of large felid ecology (body mass > 25 kg). Remote camera trap photography is the most extensively used method to identify leopards, snow leopards, jaguars, tigers, and cheetahs which feature conspicuous and easily identifiable coat patterning. Direct photographic surveys and genetic sampling are commonly used for species that do not feature easily identifiable coat patterning such as lions. We also discuss the accompanying challenges encountered in several field studies, best practices that can help increase the precision and accuracy of identification and provide generalised ratings for the common survey methods used for individual identification.
... This can be reduced by correct and protective application methods and also with the use of appropriate PPE (Damalas and Koutroubas 2016). In several occupations use of agricultural pesticides to domestic pest control has resulted in accidental poisoning (Balme et al. 2010). School-aged children and old aged population is more vulnerable to household pesticide exposure (Liu and Schelar 2012). ...
Nowadays the use of agrochemicals for agricultural farming had become inevitable towards catering the growing demand for agricultural products. Since, from the origin of green revolution the production, marketing, and use of agrochemicals had increased several folds. Only in the later years of the nineteenth century, the adverse impact of agrochemicals on the environment and human health came into light. On realizing the detrimental impacts of the overuse of agrochemicals, efforts were made to assess its toxicity to the environmental components including humans. Under this preview, this chapter describes the source and global distribution of agrochemicals in terms of its production and consumption across the world. The impact of these agrochemicals with special reference to pesticides were reviewed through their bioaccumulation and biomagnification of its residues across different trophic levels of the agroecosystem affecting the major environmental components such as air, water, and soil. Besides the residual effects on the non-targeted organisms like earthworms, fishes, birds, and humans were discussed. Humans the top predators of the agroecosystems are the worst affected due to the improper and indiscriminate use of agrochemicals, especially pesticides. Under this context exposure of humans to agrochemicals through the environment, occupation, and unexpected accidents were discussed to understand its impact on human health. Besides, the acute, sub-acute, and chronic toxic effects of pesticides were reviewed based on the recent studies conducted across the world. To understand the impact of agrochemicals on the environment, the conduct of bio-monitoring study becomes imperative, through which the risks posed by the chemicals can be assessed. The principle of bio-monitoring studies along with risk assessment and management strategies and enlisting of ecotoxicological databases were provided to better understand the adverse impact of agrochemicals on the environment and human health with further perspectives.
... Our results suggest that trophy hunting can have both positive and negative impacts: well-protected hunting areas can offer valuable habitat for leopard (Lindsey et al., 2007;Di Minin et al., 2016a;Dickman et al., 2019), but unsustainably high trophy hunting quotas may contribute to long-term leopard population declines (Packer et al., 2009(Packer et al., , 2010Loveridge et al., 2016). As such, quotas must be informed by robust estimates of population densities in hunting areas of the KAZA TFCA, ideally as part of a long-term monitoring effort with flexible quotas tied to annual population fluctuations (Balme et al., 2010a) and combined with meaningful community participation. Monitoring efforts should also be employed to identify any potential additional negative impacts of hunting on KAZA's leopard populations, such as disrupted dispersal patterns and inbreeding (Naude et al., 2020), and reduced trophy size (Muposhi et al., 2016), which can compromise the long-term sustainability of hunting activities. ...
Transfrontier conservation areas (TFCAs) have the potential to provide havens for large carnivores while preserving connectivity across wider mixed-use landscapes. However, information on the status of species in such landscapes is lacking, despite being a prerequisite for effective conservation planning. We contribute information to this gap for leopard (Panthera pardus) in Africa, where the species is facing severe range contractions, using data from transect surveys of a 30,000km² area across Botswana and Zimbabwe in the Kavango-Zambezi (KAZA) TFCA. We used occupancy models to assess how biotic, anthropogenic, and management variables influence leopard habitat use, and N-mixture models to identify variables influencing the species' relative abundance. Leopard were detected in 184 out of 413 sampling units of 64km²; accounting for imperfect detection resulted in mean detection probability p̂¯ = 0.24 (SD = 0.06) and mean probability of site use ψ̂¯ = 0.89 (SD = 0.20). Habitat use was positively influenced by prey availability and high protection. Relative abundance was best predicted by trophy hunting, which had a negative influence, while abundance was positively associated with high protection and availability of steenbok. Our findings suggest that securing prey populations should be a priority in conservation planning for leopard in Africa, and underline the necessity of preserving highly-protected areas within mixed-use landscapes as strongholds for large carnivores. Our findings also support calls for better assessment of leopard population density in trophy hunting areas, and illustrate the value of N-mixture models to identify factors influencing relative abundance of large carnivores.
... They are found in a variety of habitats, from desert to high altitude mountains and rainforests [13]. However, over the past 100 years, it is estimated that the species has declined in its historical range due to habitat degradation and fragmentation, depletion of natural prey species, poorly managed harvests, illegal trade of leopard skins and bones, and human-leopard conflict [14][15][16][17]. Leopards are killed in the Limpopo Province (South Africa) because of the threats they pose to game and livestock [18]. ...
Full-text available
The intensity and frequency of human-animal conflicts has escalated in recent decades mainly due to the exponential increase in human population in the past century and the subsequent encroachment of human activities on wilderness areas. Jhalana Forest Reserve (JFR) presents the characteristics of island biogeography in the heart of Jaipur, a city of 3.1 million people. The leopard (Panthera pardus fusca) is the top predator in this newly declared sanctuary of 29 km². We surveyed people in the 18 villages that engulf this sanctuary. We questioned the villagers’ (N=480) perceptions about conservation. Ninety-three percent (round figure) of the population have encountered leopards, and 83 % were fully aware of its role in the ecosystem. In addition, 100% stressed the necessity of conservation to save the forests and 91% supported the efforts to a wall in the reserve in order to prevent human encroachment. Most of the population is Jains and Gujars, communities which believe in non-violence. We conclude that the villagers support conservation efforts. The authorities that manage JFR view the villagers favorably and as stakeholders is the basis for continued human-leopard coexistence.
... There is clearly a need for conservation authorities to pay attention to human-wildlife conflict issues in these areas (e.g. see Balme et al., 2010). ...
... However, the sustainability of trophy hunting industry may be hinged upon several other factors such as international legislation and local governance issues (Balme et al., 2010;Peterson, 2014;Shanee, 2012); communication and branding aspects of hunting related issues (Campbell and Mackay, 2009;Damm, 2015); stakeholder and hunter perceptions, hunter attitudes and motivations (Mangun et al., 2007;Miller, 2003); hunting ethics issues (Fox and Bekoff, 2011;Lunney, 2012;Paquet and Darimont, 2010); hunting leases and hunting fees (Mozumder et al., 2007;Rhyne et al., 2009), and marketing of permits and service (Little and Berrens, 2008) among others. Although these issues are being researched nowadays, we argue that they are being done in isolation without integrating and synthesizing them for the purposes of adaptive management. ...
Full-text available
Although the contribution of trophy hunting as a conservation tool is widely recognised, there is perpetual debate and polarization on its sustainability. This review integrates five themes mostly considered in isolation, as independent research fields in wildlife conservation: (1) trophy quality and population ecology of hunted species, (2) behavioural ecology of hunted populations and associated avoidance mechanisms, (3) physiological stress in hunted populations, (4) genetic variability and desirable traits, and (5) socio-economic imperatives in wildlife conservation. We searched for articles on search engines using specific key words and found 350 articles from which 175 were used for this review under five key themes. Population and trophy quality trends of commonly hunted species seem to be declining in some countries. Elevated hunting pressure is reported to influence the flight and foraging behaviour of wildlife thus compromising fitness of hunted species. Selective harvesting through trophy hunted is attributed to the decline in desirable phenotypic traits and increased physiological stress in most hunted species. Though it provides financial resources need for conservation in some countries, trophy hunting works well in areas where animal populations are healthy and not threatened by illegal harvesting and other disturbances. There remains much polarity on the sustainability of trophy hunting in modern-day conservation. More research need to be conducted across the five themes examined in this review for broader analytical analysis and comparison purposes. A new research agenda is needed regarding wildlife sustainable use principles and their sustainability and acceptability in modern-day conservation.
... This could include faster response times from local authorities when livestock farmers perceive that they have a damage causing animal and more local government staff on the ground to verify reports of leopard predation (Chase Grey, 2011). Trophy hunting of verified problem animals could also be instituted in Limpopo Province in order to compensate livestock owners for losses caused by leopards ( Balme et al., 2010). This could improve attitudes towards leopards, foster the perception of them as economically valuable rather than as a pest species and therefore reduce retaliatory killing. ...
Human-wildlife conflict between carnivores and livestock and game owners is an issue of high conservation concern and has led to the global decline of many large carnivore species. Research has shown that carnivores are often blamed for higher levels of predation of livestock and game than actually occurs and this often leads to retaliatory killing. The aim of this study was to obtain information via scat analysis on the range of prey species taken by leopards in the Soutpansberg Mountains, South Africa, and combine these data with self-reported accounts of livestock predation from local landowners to examine differences between real and perceived leopard predation. Results showed that despite landowners reporting frequent events of leopard predation of livestock and introduced farmed game across the Soutpansberg farming community, no evidence of these species were found in leopard diets. The most frequently eaten species by relative biomass were bushbuck, hyrax and vervet monkeys; in contrast, the farmers reported cattle and impala as often being taken by leopards. Despite sharing the landscape with domestic cattle and introduced game, leopards in the Soutpansberg do not frequently utilise these species as prey and instead focus their diets on wild species. Human-carnivore conflict can be reduced by overcoming the mismatch between actual and perceived levels of predation via landowner education, effective anti-predation measures, an improved government response to reports of livestock predation and potentially giving economic value to problem animals via trophy hunting.
... An alternative to this is metapopulation management, which relies on hands-on removal and introduction of individuals between disconnected areas. Metapopulation management has demonstrated success in South Africa with African wild dogs (Gusset et al. 2006) and has been suggested as a management strategy for lions (Dolrenry et al. 2014), cheetahs (Johnson et al. 2010), and leopards (Balme et al. 2010). Expanding the current predator monitoring effort inside and outside of LBL, facilitating carnivore dispersal, and determining carnivore impacts on human communities will allow for management decisions to be made that allow for the coexistence of predators and humans alike in LBL and beyond. ...
Full-text available
The negative impact of anthropogenic activities on wildlife has led to protected areas being set aside to prevent human-wildlife conflict. These protected areas are often small and fenced in order to meet the needs of expanding human communities and to conserve wildlife. This creates challenges for the management of wide-ranging animals such as large carnivores, especially those that compete with one another for limited resources. This study focused on resource partitioning between GPS-GSM collared spotted hyenas (hereafter referred to as hyenas) and lions in Lewa Wildlife Conservancy (LWC) and Borana Conservancy (BC), Kenya. Scat analysis revealed that hyenas and lions show a high degree of dietary overlap, though hyenas have broader diets and feed on livestock species, which lions completely avoid. Spatially, hyenas show stronger intraspecific avoidance and more exclusive territorial behaviour than do lions. Hyenas and lions have a high degree of spatial overlap, though lions may influence den site selection in hyenas. Both species are heavily nocturnal and crepuscular, though hyenas tend to travel significantly further at night than do lions. Hyenas and lions both display mixed results in dynamic spatiotemporal interactions, with 40% of hyena-lion pairs showing attraction and 70% showing simultaneous use of overlapping areas. All but one inter-clan hyena pairs show strong avoidance, though lion pairs were not as mutually repulsive. This shows that hyenas and lions may use one another as sources of food and that scavenging and kleptoparasitism likely play a role in their dynamic. The hyena population is suggested to be growing and healthy, though the lion population is of concern due to lower density, isolation, and low recruitment. Further investigation into human-carnivore conflict within surrounding communities, long-term demographic and behavioural trends of all members of the large carnivore guild, and the potential development of a dispersal-based metapopulation management scheme will allow for the continued persistence of large carnivores in the Lewa-Borana Landscape (LBL) and their coexistence with human communities.
Full-text available
This study was based on a temporal analysis of trophy quality trends and hunting effort in Chewore South Safari Area (CSSA), Zimbabwe, for the period 2009-2012. We selected four of the big five species, namely; buffalo (Syncerus caffer), elephant (Loxodonta africana), the leopard (Panthera pardus) and lion (Panthera leo) for analysis. Existing database of 188 trophies from 2009 to 2011 was reviewed and recorded using the Safari Club International (SCI) scoring system. Further, 50 trophies for 2012 were measured and recorded based on the SCI scoring system. Local ecological knowledge on trophy quality and hunting effort in CSSA was obtained through semi-structured questionnaires from 22 conveniently selected professional hunters in 2012. The results indicated no significant change in trophy quality trends of buffalo, leopard and lion (p > 0.05) over the study period. In contrast, there was a significant decline in elephant trophy quality trend over the same period (p < 0.05). The results showed no significant change in hunting effort over the study period for all the four study species (p > 0.05). Furthermore, seventy-two percent (72%, n = 13) of the professional hunters confirmed that elephant population was declining in CSSA and this was likely due to poaching. Professional hunters perceived trophy hunting as a source of financial capital generation for wildlife conservation (61%, n = 11), as well as positively contributing to the local economy (56%, n = 10). It was concluded that hunting has limited negative impact on species trophy quality trends when a sustainable hunting system is consistently followed in CSSA. CSSA management need to continuously monitor trophy hunting, animal populations and employ adaptive management approach to quota setting and species conservation.
Full-text available
Humanity is exerting unprecendented pressure on natural ecosystems and the species living in them. This pressure is particularly evident among the larger members of the order Carnivora. Their large body size (typically in the 25-600 kg range), life history traits, and reliance on large prey species places them at increased risk of extinction. The International Union for the Conservation of Nature (IUCN) Cat Specialist Group, and the Convention of Migratory Species (CMS) both recognize the deficiencies in robust data available on large carnivores across large tracts of Africa. Furthermore, the population estimates we do have are often drawn from less-reliable methods. The overarching aim of this PhD thesis was to: 1) use a recently-developed population estimation technique (Elliot and Gopalaswamy 2017) to estimate the densities, population size, and population parameters of large carnivores in the Queen Elizabeth Conservation Area (QECA), Uganda, and use these data to inform their conservation status, 2) improve understanding of the conflict between large carnivores and human communities in Lake Mburo, Uganda, and Mumbai, India, and 3) explore alternative methods to fund conservation measures, including compensation and a wildlife imagery royalty. In Chapter 1 as part of introducing my thesis, I examined the literature on historic and present methods being used to census African lions Panthera leo and together with a team of international collaborators I made a case for the adoption of spatially explicit capture recapture (SECR) methods for African lions. In Chapter 2 I built upon this and showed the utility of using population state variables (namely movement, sex-ratios, and density) in assessing the conservation status of African lions in a poorly known area of East Africa. I used a population of African lions in south-western Uganda’s Queen Elizabeth Conservation Area (QECA), as a model. I conducted a 93-day African lion census in 2017-2018 and compared the results to those from an intensive radio-collaring study from a decade ago. I hypothesized that if the population of African lions in the QECA was stable or increasing, lion movement distances and home ranges would be similar between the two study periods but if movement distances were larger and sex-ratios were male-biased, the lion population was likely declining. I found male lions expanded their ranges by > 400%, and females >100%, overall lion densities were low (2.70 lions/100 km2, posterior SD=0.47), and the sex ratio of lions in the system was skewed towards males (1 female lion: 2.33 males), suggesting a decline. I concluded this chapter with a discussion of the practical conservation application of using this census technique in other parts of Africa, particularly where historic lion home-range data exist. In Chapter 3, I used the same spatially explicit capture recapture models on data collected from 74 remote camera traps set across the QECA to assess the population densities of African leopards and spotted hyenas in this savannah park. We surveyed the northern, and southern sections of the QECA, and estimated leopard densities to be 5.03 (range = 2.80–7.63), and 4.31 (range = 1.95–6.88) individuals/100 km2 respectively, while hyena densities were 13.43 and 14 individuals/100 km2. Estimates of hyena density were the highest recorded for the species anywhere within their range using SECR methods. I also suggested that the high hyena densities could be related to the evidence provided in Chapter 2 of African lion decline in the QECA. One hypothesis that could explain the inverse densities of hyenas and lions is that hyenas have experienced competitive release from African lions in the QECA. Similar findings have been reported in the Talek region of Kenya’s Maasai Mara, and Zambia’s Liuwa Plains. This chapter also provided the first SECR population estimates of leopards, and spotted hyenas anywhere in Uganda. In Chapter 4 and Chapter 5 I addressed the most important threat to the existence of large carnivores: conflict with human communities, and their livestock. While conflict tends to dominate the narrative where large carnivores and humans co-exist, there can often be direct and indirect benefits to humans. In Chapter 4 I examined the ecosystem services provided to people by the Indian leopard Panthera pardus fusca, in Mumbai.The Sanjay Gandhi National Park (SGNP) is located in the city of Mumbai, India, and has some of the highest human population densities in the world. Large carnivores are known to control prey populations, suppress smaller carnivores, reduce parasite load in humans, and promote seed dispersal. However, this chapter is one of the first studies highlighting the ecosystem services provided by a large carnivore outside of a natural or protected system. I showed that leopard predation on stray dogs reduced the number of people bitten by dogs, reduced the risk of rabies transmission, and reduced dog sterilization and management costs. Our estimates showed that dog densities around SGNP (17.3/km2) were 40 times lower than four nearby urban informal settlements (688/km2) and were ten times lower than the citywide mean (160/km2). If it is, as we propose, leopards that are holding the dog population around the park at its current density, dog bites could increase from 3.6 bites/1000 people to 15.5 bites/1,000 people if leopards were to disappear. As over 78% of dog bites in Mumbai require treatment, and 2% require rabies post-exposure vaccination, the treatment costs could reach as high as US$ 200,000 per year (compared to ~US$ 42,500 currently). As development pressures are threatening the region’s leopards, this work shows the potential costs of their local extirpation. Chapter 5 assesses the landscape-level correlates of livestock attacks by two large carnivores, the spotted hyena, and African leopard in the cattle and sheep/goat farms bordering Lake Mburo National Park, south-western Uganda. I also make suggestions on how to improve the sustainability of a voluntary financial compensation scheme run by a local lodge (the Mihingo Conservation Fund) aimed at alleviating persecution of these species. I used ten years of depredation events to investigate the importance of seasonality and landscape features (ie. terrain ruggedness, proximity to roads, water, human settlements, and vegetation density) on livestock attack probability. I also examined the current costs of the compensation scheme of reported attacks. I showed that most livestock attacks in this region were caused by spotted hyenas, both predators killed at night, did not exhibit seasonal patterns in depredation, and attacks were owed to poorly fortified bomas (82% of leopard attacks and 64% of hyena attacks were made inside bomas). Attacks were also made near human settlements, close to the national park border, and in areas of rugged terrain. The compensation fund made more gross income from tourism activities than was paid in compensation in most years, but compensation costs had to be subsidised by the lodge because the funding was also used in other community development projects (eg. building of a school, and paying children’s school fees). Chapter 6 of this thesis built upon the sub-theme of Chapter 5, funding of carnivore conservation measures and created a roadmap for a recently proposed idea of a threatened wildlife imagery royalty to stem the large budgetary shortfalls facing large carnivore conservation. The idea of a threatened species imagery royalty was proposed in two recent papers, Good et al. (2017) and Courchamp et al. (2018). I built upon these and discussed how such a royalty could be implemented, explored several legal avenues for its application, and also showed its potential scale in leveraging funding. The creation of a national law which charges a royalty from corporations using the imagery of their threatened wildlife, and a “Fairtrade” equivalent held the most promise for the development of a wildlife imagery royalty. Indeed, articles 3 and 6 of the Convention on Biological Diversity (CBD) encourage sovereign states to ensure activities within their jurisdiction and control do not damage the environment of other states. Similarly they are encouraged to develop national strategies, plans or programs for the conservation and sustainable use of biological diversity. The funding that could potentially be leveraged from a wildlife imagery royalty is immense. I used large felids as a model group to show that the relevant 14 companies on the Forbes 2000 list alone could generate US$ 202 million–2.02 billion if they paid 0.1-1% of their profits in royalties. My thesis addressed an important but often overlooked component of estimating large carnivore populations, the use of population state variables in informing conservation status. The use of animal movement, sex-ratio, and density information has wide application that transcends large carnivores. My assessment of leopard-dog interactions, and the potential implications for humans, was one of the first examples in the literature of the potential benefits a large carnivore may have to humans. The assessments of compensation and wildlife imagery royalties have important consequences on better managing and also leveraging funding for the conservation of large carnivores and other threatened, enigmatic species.
ResearchGate has not been able to resolve any references for this publication.