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Managing Charismatic Carnivores in Small Areas: Large Felids in South Africa


Abstract and Figures

Large carnivores are key foci for conservationists, tour operators and hunters alike. They provide revenue-generating opportunities, but also can be keystone species in conservation areas, influencing the maintenance of biological diversity. They often degrade livelihoods of people when coming into conflict with livestock land-uses. We acknowledge these challenges specifically for cases where large carnivores are present in small areas and propose an alternative strategy to the traditional carrying capacity approaches, directed at managing the effects of large carnivores. We advocate an approach where managers of small areas mimic natural social dynamics such as coalition tenure, density dependent changes in litter size, age at first birth and birth intervals, as well as subadult dispersal. This assists with achievement of population and evolutionary targets through a process-based approach mimicking drivers of variance in social groups. Such an applied conservation husbandry approach may have robust outcomes that do not compromise conservation values.
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Managing charismatic carnivores in
small areas: large felids in South Africa
Sam M. Ferreira1*& Markus Hofmeyr2
1Scientific Services, SANParks, P.O. Box 202, Skukuza, 1350 South Africa
2Veterinary Wildlife Services, SANParks, Skukuza, 1350, South Africa
Received 2 January 2013. Accepted 3 September 2013
Large carnivores are key foci for conservationists, tour operators and hunters alike. They
provide revenue-generating opportunities, but also can be keystone species in conservation
areas, influencing the maintenance of biological diversity. They often degrade livelihoods
of people when coming into conflict with livestock land-uses. We acknowledge these
challenges specifically for cases where large carnivores are present in small areas and
propose an alternative strategy to the traditional carrying capacity approaches, directed at
managing the effects of large carnivores.We advocate an approach where managers of small
areas mimic natural social dynamics such as coalition tenure, density dependent changes in
litter size, age at first birth and birth intervals, as well as subadult dispersal.This assists with
achievement of population and evolutionary targets through a process-based approach
mimicking drivers of variance in social groups. Such an applied conservation husbandry
approach may have robust outcomes that do not compromise conservation values.
Key words: African lion, cheetah, small reserves, social dynamics, conservation husbandry.
The restoration and maintenance of predation is
often a key aim for conservation agencies wishing
to achieve ecosystem objectives (Dalerum
et al.
2008). The challenges of maintaining ecological
processes that involve large carnivores are com-
plicated by expectations and attitudes of stake-
holders (Kerley
et al.
2003). These range from
positive, such as a general assumption that the
presence of large carnivores enhances a tourism
experience (Okello
et al.
2008), to negative (Treves
& Karanth 2003), such as expectations and atti-
tudes towards livestock losses as a result of carni-
vores spilling over into landscapes abutting
protected areas (Inskip & Zimmermann 2009). In
some instances, populations of large carnivores
may have ranges largely outside protected areas
et al.
The ecosystem management paradigm (Bocking
1994), adhered to in some instances by conserva-
tion agencies (Rogers 2003), recognizes ecologi-
cal processes as key drivers of how the larger
mammalian herbivores influence plant community
structure and function in ecosystems (Gordon
et al.
2004). This implies top-down regulation of
plant community structure and function. However,
it is likely that both top-down, through predation,
and bottom-up, through nutrient quality of vegeta-
tion, may regulate herbivore dynamics (Turkington
2009). The absence of large predators may thus
pose some constraints on conservationists wish-
ing to restore key ecological processes associated
with herbivores and how predation influences
herbivorous effects.
In South Africa, lions (
Panthera leo
) and cheetahs
Acynonyx jubatus
) have disappeared from most
of their historical range (Nowell & Jackson 1996;
et al.
2007). Lion populations survive in
Kruger National Park, Kgalagadi Transfrontier
Park and the Greater Mapungubwe Transfrontier
Conservation Area including farmland (Nowell &
Jackson 1996). Cheetahs also occur in these pro-
tected areas and associated surrounds as well as
on farmland across the North West and Limpopo
provinces of South Africa (Marnewick
et al.
Reintroductions of both these species (Hayward
et al.
2007; Hunter
et al.
2007; Marnewick
et al.
2007; Funston 2008; Davies-Mostert
et al.
Slotow & Hunter 2009) since the early 1990s into
small, fenced areas including different types of
administrative and legislative land-use designa-
tions such as private reserves, conservancies,
protected areas such as TFCAs, national and pro-
vincial parks, have resulted in several free-ranging
populations in small areas (<1000 km2) isolated
from each other.
Recovery of ecosystems following disturbances
is not haphazard and species usually colonize
South African Journal of Wildlife Research 44(1): 00–00 (April 2014)
*To whom correspondence shoud be adressed.
degraded or disturbed areas in predictable
sequences (Wassenaar
et al.
2005). Restoration
of carnivore assemblages should therefore also
not be haphazard. Succession theory predicts
that smaller generalist herbivores will colonize a
disturbed area first, followed by larger herbivores
et al.
2001). The relevant predators
specializing on each herbivore assemblage (see
Owen-Smith & Mills 2008) should follow a similar
trend. Active carnivore restoration, like that prac-
ticed in South Africa (Hayward
et al.
2007; Hunter
et al.
2007; Marnewick
et al.
2007; Funston 2008;
et al.
2009; Slotow & Hunter
2009), may thus better succeed in establishing
dynamic predator–prey relations if such a coloni-
zation sequence is mimicked through reintro-
ductions (Sait
et al.
2000). Smaller carnivores
should thus be reintroduced first.
An additional challenge most pertinent in small
reserves is the consequences for ecological
(Hayward & Kerley 2009) and genetic (Trinkel
et al.
2008; Trinkel
et al.
2010; Trinkel
et al.
2011) integ-
rity of populations of large carnivores and prey
alike. Most carnivore species will breed success-
fully if released into a fenced reserve with high
densities of predator-naïve prey (Hayward
et al.
2007). In addition, confinement influences where
carnivores and prey can move, which can alter
the way they interact (van Dyk & Slotow 2004;
Hayward & Kerley 2009). Furthermore, their influ-
ences on ecosystems as a result of their top-down
regulating roles in ecosystems (Owen-Smith &
Mills 2008) can deplete the prey base in a non-sus-
tainable way in spatially limited environments
(Slotow & Hunter 2009). In some instances regular
prey introductions are necessary to sustain a
carnivore presence (
et al.
2009). Even though the response to a large carni-
vore introduction may be seen as a compensatory
drop in prey abundance, spatially restricted and
fenced environments limit the recovery of prey
species as no opportunity for immigration to take
place exists unless mimicked artificially (Hayward
& Kerley 2009). Consequently, restricted areas
may often have limited spatial and temporal varia-
tion in predation risk, vulnerability and fear land-
scapes. These then lead to a disruption of dynamic
predator–prey relationships that may undermine
ecological objectives.
Small spatially restricted or fenced areas may
also experience meso-predator releases because
of the absence of large carnivores. Competitive
interactions with large carnivores (Ritchie & John-
son 2009) may limit spatial and temporal dynamics
of the smaller carnivores. This has two conse-
quences when large predators are absent, meso-
predator release may result in spill-over effects
that accentuate human-wildlife conflict in areas
abutting small restricted areas (Prugh
et al.
Secondly, upon introduction of larger carnivores,
persistence of rarer small carnivores may be
challenged through meso-predator suppression
(Ritchie & Johnson 2009).
In addition to the above ecological challenges,
charismatic carnivore reintroductions are often
token ecological actions because the primary
motive is the accrual of benefits from large carni-
vore eco-tourism (Kerley
et al.
2003, Okello
et al.
2008). This is defendable for privately owned
areas, but is questionable when introductions
directed at improving a tourism product take place
in protected areas. The National Environmental
Protected Areas Act (Act No. 57 of 2003) of South
Africa provides for such activities as long as the
ecological integrity of the protected area is not
jeopardized. The disruption of prey-related pro-
cesses when confined in small areas may be
defined as illegal and in contravention of the act.
Managers of protected areas are thus particularly
accountable for dealing with disrupted predator
prey relations when large carnivores have been
introduced into small areas.
These observations and constraints have led to
carnivore management approaches for small
areas that define some or other ‘carrying capacity’
of predators based on available prey (Lindsey
et al.
2004; Hayward
et al.
2007; Lindsey
et al.
However, developments within the field of wildlife
management have seen the adoption of a conser-
vation paradigm that acknowledges the flux of
nature and importance of spatio-temporal patchi-
ness and habitat heterogeneity (Stalmans
et al.
2001). Such patchiness should also be reflected in
predation risk, vulnerability and fear landscapes.
Consequently the shift away from the ‘carrying
capacity’ concept in conservation management
et al.
2009) to one that considers the
dynamic equilibrium and change inherent within
such complex systems (Wu & Loucks 1995) has
been adopted by some conservation agencies
Rogers 2003) in order to manage ecosystems
to meet biodiversity and other conservation objec-
Here we provide an alternative conceptual model
to the carrying capacity approaches adhered to at
present (Lindsey
et al.
2004; Hayward
et al.
2 South African Journal of Wildlife Research Vol. 44, No. 1, April 2014
et al.
2011). Our approach hinges on
mimicking natural social dynamics. We do not
compare carrying capacity approaches with the
proposed alternative primarily because our
approach is embedded within a system of flux
management philosophy (Rogers 2003). Carrying
capacity approaches imposes non-flux philoso-
phy. We use lions (
Panthera leo
) and cheetahs
Acinonyx jubatus
) to illustrate the potential appli-
cation and discuss the consequences and pitfalls
that social management of carnivores may pose.
Large carnivore abundances and densities are
associated with the quantity and spatial distribution
of prey biomass (van Orsdol
et al.
1985, Packer
et al.
2005, Hayward
et al.
2007). This relationship
is influenced by the variable social nature of most
large carnivores (
Bertram 1973). Social factors
may limit minimum home range sizes and often
reduce survival and fecundity. Free ranging carni-
vore populations in large spatially non-restricted
areas are often variable and fluctuate in abun-
dance, but seldom are long-term trends positive or
negative (Packer
et al.
2005, but see Vucetitch &
Peterson 2004).One consequence of fenced con-
fined areas is an increase in the likelihood that
individuals may socially become more tolerant of
each other as is the case when unrelated lions are
placed in a small boma (M. Hofmeyr, unpubl. data).
In small areas spatial restrictions may thus remove
one of the key mechanisms of population regula-
tion of large carnivores,
social unfamiliarity.
Conceptually, management of large carnivores
in spatially restricted areas could focus on manag-
ing social relations, which is one of the key mecha-
nisms of carnivore population regulation (Bertram
1973). This may ultimately also drive variability in
predation risk, vulnerability and fear landscapes,
rather than a population approach with fixed
‘carrying capacity’ as was historically the case
et al.
2007). Given this context, most
small protected areas and private reserves in
South Africa are only capable of dealing with one
or two social groups of a particular species of large
carnivores in the various management units.
Carnivore management in restricted areas may
achieve better outcomes by focusing on social
units rather than managing a carnivore population.
Solitary and small carnivores typically provide
fewer management challenges.
Managing social relations contrasts starkly with
the population approaches typically adopted for
large herbivores (Gordon
et al.
2004). Social fac-
tors may significantly be important in regulating dy-
namics through effects on survival and fecundity
(Funston 2001). Their resource needs necessitate
large individual home ranges, which are often
vigorously defended, and makes them more sus-
ceptible to spatial restriction (
et al.
2005). They have rapid impacts on their resources
and other smaller carnivores (Ritchie & Johnson
2009), but resources can also recover rapidly
et al.
2010). They have short life histories
(Gittleman 1986), which mean that they have
inherent demographic vigour to respond fast to
management interventions. And finally, managing
carnivore social units by mimicking social dynamics
focuses on the biological mechanisms (see
et al.
2011) that are most likely disrupting
predator–prey relationships and reducing genetic
Our conceptual model thus suggests that con-
servationists mimic behaviour and dynamics of a
social group. This means mimicking sex-specific
dispersal events, dominance hierarchies, socially
induced reductions in birth rates, socially induced
mortality events and variance in these variables.
Each of these varies in free-ranging large carni-
vore populations. Such variability should be part
of the mimicking of social dynamics in spatially
restricted areas and is likely to produce heteroge-
neity in group sizes as well as temporal and spatial
predation pressures that are more likely to induce
persistent dynamic predator–prey relations.
Managing lions
Female lions conceive as early as 32 months
(Hunter 1998) and given their gestation of 14–15
weeks may have their first litter when between
40 and 60 months old (Smuts
et al.
1978). In small
confined populations lions tend to breed at younger
ages (Hunter 1998). In large free-ranging popula-
tions litter sizes vary from one to six (Rudnai 1973;
Packer & Pusey 1987; Patterson 1988), but gener-
ally converges onto an average of three (Smuts
et al.
1978; Hunter 1998).
Birth intervals depend on whether cubs are
raised to maturity. New litters are born every 20 to
24 months (Packer & Pusey 1987; Hunter 1998).
When litters are lost, intervals range between four
and six months (Packer & Pusey 1987).
Cub mortality is high (Orford
et al.
1988). How-
ever, in environments like Kruger National Park, up
to 80% of cubs observed survived the first year
(P. Funston, pers. obs.). Cub mortality increases
Ferreira & Hofmeyr: Managing charismatic carnivores in small areas 3
when male coalitions take over tenure of a pride
et al.
2008). Subadults and adults have
higher survival rates than cubs based on data
compared between Kruger demographic surveys
(Ferreira & Funston 2010) and long-term studies in
the Serengeti (Packer
et al.
To mimic social dynamics for females, five
management options are available:1) increase the
age at first reproduction using contraception of
subadult females; or 2) establish longer intervals
between births using contraception of adult
females (Munson 2006; Bertschinger
et al.
3) reduce age specific fecundity through reduced
litter sizes using unilateral tube-tying of fallopian
tubes (Alhasani
et al.
1984); 4) mimic female
dispersal by removing as well as introducing
subadult females; 5) mimic higher death rates of
old females by removing the oldest females in the
pride. Unilateral tube-tying of fallopian tubes may
require experimental evaluation and clinical
improvement before implementation. A sensible
target is the maintenance of variance in the
number of adult females comprising a pride
centring on that observed in unrestricted free-
ranging populations. This converges onto four
although a range of two to 11 has been noted in
several places (Schaller 1972a; Smuts 1976;
Stander 1991; Funston 2001).Managers may thus
use modelling approaches (Quadling & Starfield
2002) to define a mixture of the five options to
achieve an average pride size of four.
Male survival rates are typically lower than those
of females (Ferreira & Funston 2010), but this
discrepancy dissipates when in small confined
areas (Slotow & Hunter 2009). Three manage-
ment options are available for males: 1) Mimic
male dispersal through removal and introductions
of subadults; 2) mimic pride take-overs – tenure is
on average three years (Funston
et al.
and 3) mimic higher death rates of old males by
removing the oldest males. Managers can again
use modelling approaches (Whitman
et al.
et al.
2007) to achieve at least coalition
tenure of three years and allowing each coalition to
have only one breeding opportunity. No active
switches of coalitions may be required if subadult
male introduction and the oldest male removals
are staggered. A pride take-over should then follow
naturally, with subsequent consequences for cub
mortality.Tolerance of older cubs may be possible
(M. Hofmeyr, pers. obs.) so introduction of new
males could be planned at times when females
have young cubs.
Managing cheetahs
Cheetahs are generally solitary except for male
coalitions and females with dependent cubs (Caro
1994). Social structure is regulated by males
marking their territory and patrolling it, defending it
aggressively against other males (Caro & Collins
1987a). Males may also fight to the death over
females (Caro & Collins 1987b, M.G.L.Mills, pers.
comm.) By contrast, females do not defend the
ranges they occupy (Caro 1994).
Female cheetahs typically conceive for the first
time at age 21–28 months (Labuschagne 1979;
Pettifer 1981; Kelly
et al.
1998) with a gestation of
90–98 days (Schaller 1972b,c,d). They may thus
have their first litter when 24–31 months old.They
tend to have litters once every 18–20 months,
although should they lose a litter earlier they will
quickly breed again (Schaller 1972b,c,d; Frame &
Frame 1976;Kelly
et al.
1998) and litter sizes range
from two to six (Schaller 1972b,c,d: Frame &
Frame 1976; Pettifer 1981; Hunter 1998).
Cubs reach independence of females at
13–24 months (Schaller 1972b,c,d). After leaving
the mother the sibling group usually stays together
for at least a few months (Caro 1994.), even up to a
year (M.G.L. Mills, pers. obs.). Females typically
leave the sibling group on their own when they
come into estrus (Caro 1994; M.G.L. Mills, pers.
obs.), while male litter mates form coalitions of,
usually two to three, but up to four to five individuals
(Caro & Collins 1987b). Males generally only join
with a female when she is in oestrus (Hunter
1998). Cub survival rates are variable, but can
range from 5% (Laurenson 1994) to 36% (Mills &
Mills, in press). Where there are many competing
predators, especially lions, spotted hyaenas and
leopard, cheetah live at low densities and have low
survival rates. In fenced parks, where there are no
other large predators, and areas with low large
carnivore density, sur vival rates are higher (62% of
those that had left the den in Phinda Game Reserve,
Hunter 1998, and 64% in the Kgalagadi Trans-
frontier Park – Mills & Mills, in press) and females
have raised as many as six cubs successfully
(Hunter 1998).
The effect of social dynamics on female cheetahs
can be mimicked through the same five manage-
ment options outlined for lions. Typically, two to
eight females will share parts of their home range
et al.
2008). A sensible target is the main-
tenance of variance in the number of adult females
that may frequent management units. Managers
can make use of modelling approaches (Crooks
4 South African Journal of Wildlife Research Vol. 44, No. 1, April 2014
et al.
1998; Lubben
et al.
2008) using a mixture of
options to achieve an average number of adult
females of four in each management unit.
Male survival may be lower than that of females
because they aggressively defend territories (Hunter
& Skinner 1995). When male cubs become inde-
pendent, they disperse with their sibling brothers
until mature enough to compete with other male
coalitions. Single males may join other males in
coalitions, or remain solitary. Territorial tenure of
male coalitions can last from three to 25 months
depending on the size of the coalitions. Occa-
sionally tenure may last for an extended period up
to six years (Caro & Collins 1987a; Caro & Collins
Three feasible management options are avail-
able for males: 1) mimic male dispersal through
removal and introductions of subadults; 2) mimic
territory tenure, and 3) mimic higher death rates of
old males by removing the oldest males. Territory
sizes range from 37 km2to 504 km2(Pettifer 1981;
Caro 1994; Hunter 1998; Mills 1998). Given the
range of sizes in the management units compris-
ing the typical reserve, a single coalition with the
occasional presence of two coalitions may be a
sensible target. Alternatively, single males may be
just as successful. Using modelling approaches
et al.
1998; Lubben
et al.
2008) managers
can design an intervention programme to achieve
territory tenure of two years and allowing each
coalition to have only one breeding opportunity.
Managing cheetah at this scale is much more diffi-
cult than lions and will need most cheetahs to be
collared to facilitate efficient monitoring.
Managing large carnivores in small areas carries
several challenges. We suggest that social
mimicking acting as an alternative to carrying
capacity approaches (Hayward
et al.
2007) should
lead to variable population sizes that may be more
compatible with system-based approaches to
conservation management (Rogers 2003). Our
approach hinges on the assumption that social
limitation plays a key role in regulating the dynam-
ics of social (
lions, Bertram 1973) or even
asocial carnivores living in groups (
This implies that on a local scale when restricted
space interferes with social dynamics, predator–
prey dynamics revert to numerical drivers rather
than spatial as has been proposed for some
mega-herbivores (van Aarde
et al.
2006). How
sensible is this proposition?
We argue that systems-based approaches to
conservation management (Bocking 1994) have
three elements; providing opportunities for ecolog-
ical processes to play out, restoring ecological
processes and/or mimicking the outcomes of
ecological processes. The four key population
processes birth, deaths, immigrations and emigra-
tions are what our suggestion targets, but in a
mechanistic way (Ferreira
et al.
2011) at the social
group level.Is there biological reality to mimicking
these social effects on vital rates?
Lions provide an example of the mechanistic
breakdown of social systems in small confined
areas. Female lions live in groups to either defend
cubs from roaming males or defend territories from
other groups (Packer
et al.
2005). Defense is asso-
ciated with lion-lion interactions that may induce
stress and carry risk for individual survival. In small
confined areas roaming males are often absent
and these areas more often than not only contain
one pride. The drivers of group living are dimin-
ished. It is predicted that mortality risks and social
stress weakens in such cases.
Females may lose body condition when socially
or resource stressed (Clauss
et al.
2010). Young
females that have never bred may not have good
enough body condition to enter reproductive activ-
ity. This mechanism thus increases age at first
birth. Carnivore females that have already bred do
not reach good enough condition to become recep-
tive and do not conceive. This typically increases
birth intervals. Alternatively, females that have
already conceived suffer from maintaining good
enough breeding condition and abort or resorb
fetuses (
Greenwood & Sargeant 1994) lead-
ing to reduced litter sizes. Given that resource
stress and social stress are often expected to be
greatly reduced in small areas, we propose that
the mechanisms that reduce fecundity are absent
in such areas. Our propositions focus on inducing
such control on fecundity by mimicking the out-
comes of social stressors and thus carry biological
Survival effects also seem to be associated with
resources as well as social stresses.For both lions
and cheetahs, survival is associated with prey
biomass, but social interactions irrespective of
resources also greatly influence survival (Durant
et al.
et al.
2010). Coalition take-over
for instance is a key driver of lion cub mortality
et al.
2008) as well as subadult male
survival in cheetahs (Durant
et al.
2004). Such
mortalities may in turn be a key driver of carnivore
Ferreira & Hofmeyr: Managing charismatic carnivores in small areas 5
dynamics or at least a key determinant of variance
in carnivore dynamics both spatially and tempo-
rally (
Funston 2008). That in turn should affect
predator–prey relations simply through resultant
variance in carnivore abundance.
The relative role of fecundity versus cub survival
in affecting recruitment of individuals into the
breeding adult populations of both species is
largely unknown. We anticipate that this may differ
from case to case and could depend on a variety of
factors including environmental and social. Even
so, a significant criticism of our approach could
focus on why our proposed actions do not actively
induce cub mortalities.
In terms of our social management framework,
the mimicking of variance in cub survival is a
by-product in our mimicking of coalition take-over
in lions, which is one of the key mechanisms induc-
ing variance in cub-survival (Lehmann
et al.
The lack of resources leading to nutrition related
mortalities when intra-specific competition is
strong (
Sliwa 2006) is a dynamic driver of
carnivores related to the density of prey (
Ferreira & Funston 2010). In small areas where
accessibility to prey may be higher irrespective of
prey density (Funston 2008), the above mecha-
nism is degraded and cub survival tends to be
higher. Our approach targets part of the nutrition
mechanisms through the spin-offs provided.
Secondly, the euthanasia of cubs may be ethi-
cally unacceptable to society at large (Fahrion
et al.
2011). We do not exclude cub euthanasia as
in some cases it may be the only practical solution
to mimic some of the natural processes that
free-ranging lions and cheetahs in large areas are
exposed to.However, by focusing on the fecundity
component of the recruitment dynamics of adults,
a much more ethically acceptable option is pre-
ferred as part of achieving a variety of socially
responsible conservation mandates.
A key concern is that culling and husbandry type
approaches impose large and rapid selection
pressures on a population (
et al.
2001). This could have far reaching consequences
for evolutionary potential and directly contrast eco-
system-based approaches (Bocking 1994). Our
approach of allowing coalitions essentially one
opportunity to breed and having external linkages
in the network of management units is likely to
maintain evolutionary potential as well as genetic
diversity over time. We reason that given that
social dynamics similar to carnivores in large
areas are mimicked, genetic integrity will be an
outcome, in stark contrast to approaches used
until now (see Trinkel
et al.
We have argued strongly for a social approach to
mimicking social dynamics for group or near-group
living carnivores when these live in spatially
restricted areas. This suggestion, however, carries
some pitfalls, most of which are speculative. First,
anecdotal cases have been noted where contra-
ception imposes individual health threats. Health
complications may arise in later reproductive
physiological responses when females are chemi-
cally prevented from having their first cycle. In
addition, health complications may arise when
females that have been on contraceptives for
extended periods are allowed to cycle again,
but do so irregularly (
et al.
Furthermore, the physical tying of uteri tubes
(unilateral salpingectomy, Alhasani
et al.
1984) to
reduce litter size may lead to anatomical complica-
tions, surgical complications and associated infec-
tions. This reflects some of the concerns associated
with the application of contraceptives to other
carnivores in free-ranging environments (Grandy
& Rutberg 2002).
Secondly, at a behavioral level, change in the
demand of dependents on adults may induce
behavioural changes (
et al.
2002). For
instance, if an individual female’s resource need is
substantially reduced it may allow individual hunt-
ing and lead to fragmentation of prides. This is
likely to conflict with some of the systems-based
conservation objectives such as providing for
tourism experiences, and increasing overall pre-
dation rates (Okello
et al.
Thirdly, although our proposition implies popula-
tion-level responses as potentially desirable, the
consequences for predator prey-interactions are
not clear.The spatial distribution of resources and
new needs may impose different home range and
spatial use patterns of manipulated individuals
Kerley & Shrader 2007), which in turn may
disrupt predator–prey interactions across land-
scapes. We propose that these concerns and
speculations, structured as hypotheses, should
form a key part of evaluating the consequences of
a socially-based approach to carnivore manage-
ment. Evaluation should focus on consequences
at the individual health, behavior, population, evo-
lutionary and predator–prey interaction level.
The implementation of our strategy may be seen
as equivalent to animal husbandry (
et al.
2006) or captive breeding (
Williams &
Hoffman 2009) approach, but the outcomes are
6 South African Journal of Wildlife Research Vol. 44, No. 1, April 2014
vastly different. Our suggestions do not focus on
maximizing productivity (Shipilov 1972) as is
the case for husbandry and captive breeding
approaches, but rather inducing variance in dynam-
ics by mimicking the mechanisms that would do
that in free-ranging populations. The approach is
intensive and potentially costly and requires many
linkages and management agreement between
several conservation areas. The number of link-
ages depends on species life-histories as well as
the number and size of management units.
Our approach carries potential implications for
carnivore regional conservation plans. For one,
integration may be more successful with such a
process-based approach, but it may challenge
objectives of meta-populations, in particular for
cheetah meta-populations (Johnson
et al.
because our approach essentially recognizes a
single population with social groups spatially iso-
lated over vast areas.That is because meta-popu-
lation theory, based on asynchronous subpopulation
dynamics, variable dispersal and availability of
empty habitats (Olivier
et al.
2009) makes sub-
stantially different predictions than those made by
single population paradigms.
Some of the above challenges can be reconciled
through identifying regional nodes (
Fig. 1 for
lions and see Marnewick
et al.
2007 for cheetahs)
based on genetic geographical structure. We
suggest that most integration is then within such
nodes with rare external linkages to other nodes.
The reality is that regional plans may benefit
administratively, logistically, but most importantly,
ecologically from such an integrated process-
based husbandry approach. We caution though
that although our approach hinges on the assump-
tion that social limitations play key roles in regulat-
ing dynamics of social (
Bertram 1973), or even
asocial carnivores living in groups (
ecological conditions may also affect social behavior.
This suggest challenges for a blanket policy for all
reserves in all habitats, some of which may be
Ferreira & Hofmeyr: Managing charismatic carnivores in small areas 7
Fig. 1.Lion case study serving as an example of potential regional nodes for social-based large carnivore manage-
ment in South Africa. Four populations comprising spatially isolated social groups (¡) combine to form a single
meta-population across South Arica.
overcome by defining regional nodes based
on broad biome characteristics. Even so, we
acknowledge that a process-based or conserva-
tion husbandry approach requires buy-in and
coordination, but advocate that it provides a viable
alternative to tourism driven management inter-
ventions and can create dynamic predator–prey
relations in small areas addressing a variety of
conservation objectives.
Conservation implications
We have suggested an alternative approach for
large carnivore management in small areas, some
of which do not have biodiversity objectives as key
foci. The approach suggests that site- and case-
specific population and evolutionary targets can
easily be met through a process-based approach
that attempts to mimic the various mechanisms
that drive variance in social groups. This is a
unique and contrasting way to apply husbandry
approaches and requires rigorous evaluation at
several levels. Even so, the approach suggested
here may produce significant outcomes that are
more defendable and acceptable to scientists,
managers and the general public at large without
compromising biological and other conservation
This manuscript benefitted from discussions
with Park Management staff and the regional
ecologists of the Arid, Frontier and Northern
Regions of parks under the management authority
of SANParks.
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Ferreira & Hofmeyr: Managing charismatic carnivores in small areas 11
... In addition, lioness reproductive success declined, and injury level increased, with an increasing number of adult female neighbours ). Furthermore, lionesses from smaller prides that were surrounded by more female neighbours were less often alone than those in smaller prides with fewer neighbours (Packer et al., 1990 (Ferreira & Hofmeyr, 2014). In addition, solitary lions are known to be effective hunters (Caraco & Wolf, 1975). ...
... In addition, solitary lions are known to be effective hunters (Caraco & Wolf, 1975). Thus, reduced cohesion within prides can increase predation pressure and degrade natural predator-prey dynamics in fenced reserves because there are more individuals potentially hunting more prey (Ferreira & Hofmeyr, 2014). Furthermore, prey abundances generally remain high ...
... Mimicking meta-population dynamics by facilitating lion social processes such as dispersal and male tenure turnover (Ferreira & Hofmeyr, 2014) is embedded within a suite of best practice guidelines adopted by managers of fenced reserves (Miller et al., 2013). Adoption of such interventions which likely induce meta-population dynamics, however, is a recent phenomenon and managers must routinely address the consequences of past, haphazard management interventions (Miller et al., 2015). ...
Full-text available
African lion (Panthera leo) populations normally consist of several neighbouring prides and multiple adult males or groups of males that interact competitively. In large, open systems, cub defence from infanticidal males and territory defence drive group living in lions. However, in smaller (<1000 km2), fenced wildlife reserves, opportunities for natural immigration and emigration are limited which means that the evolutionary drivers of lion sociality may collapse. Here, we use lion behavioural data collected from 16 wildlife reserves across South Africa to test how management‐induced ecological conditions alter lion social dynamics. The number of lionesses observed together was best predicted by pride size, prey biomass and biome. Lionesses were less likely to group together as pride size increased, but more likely to group together as prey biomass and habitat productivity increased. In addition, adult males were observed more frequently with prides that had young (<12 months) cubs in reserves that had unfamiliar adult males present compared to reserves without any unfamiliar adult males. Our results demonstrate how intraspecific competition between lions drives their sociality, and this may break down in small, fenced wildlife reserves where lions are actively managed. Although small, fenced reserves in South Africa have made a significant contribution to increasing lion numbers on the continent, our work highlights several important ecological implications of active lion management. For wildlife managers, mimicking the outcomes of different levels of intraspecific competition is likely a critical management tool for the persistence of lions in small reserves. The number of lionesses sighted together relative to pride size varied across four categories of lion populations in South Africa (A–D). The probability of observing a separate pride increased with increasing pride size, except for managed wild prides in reserves with more than one pride (category B).
... This management approach can reduce the need for lions to forage outside of their normal territories and risk encountering extra-pride groups (Mosser, Fryxell, Eberly & Packer, 2009;). Furthermore, lioness body condition, which affects reproductive rate, will likely be higher in small reserves that have higher prey biomass (Ferreira & Hofmeyr, 2014). ...
... Lion populations persisted at higher densities in the Nairobi NP (Rudnai, 1973), comparable to the maximum densities recorded in our study, likely associated with the proximity of Nairobi to the equator (sensu Willig, Kaufman & Stevens, 2003). The Nairobi lion population also persisted at high prey availability and lion emigration rates (Rudnai, 1973), both of which managers unintentionally mimicked in small, fenced reserves (Ferreira & Hofmeyr, 2014) and likely further increased lioness reproductive parameters. The lion populations in Nairobi NP as well as our study had high reproductive rates, indicating that they did not exceed optimum densities (Rudnai, 1973). ...
... Therefore, it is possible that lionesses in our study also mated with the small number of extra-pride males (when present) during oestrus. Lioness avoidance of extra-pride males was therefore possibly lower in our study and may have contributed to the younger age at first birth in conjunction with increased body condition as a result of reduced resource competition (Ferreira & Hofmeyr, 2014). Without evidence of extra-pride copulations in small reserves, future genetic work should investigate this possibility, along with management records of any such instances. ...
... This highlights two important considerations for managing highly social species. Firstly, the number of groups may be more important than the number of individuals (Ferreira and Hofmeyr 2014), and, secondly, social groups are fundamental to (Konstant and Mittermeier 1982) and fossorial European ground squirrels (Spermophilus citellus; Gedeon et al. 2011). While shelter types tend to be species-specifi c, this management action again leads to the potential creation of ecological imbalances favoring individual species at the expenses of others. ...
... In this case, young elephants that were orphaned during culling operations in Kruger National Park were reintroduced to Pilanesberg, and as they became sexually mature, started killing black and white rhinoceros . The introduction of older humanity (see Ferreira and Hofmeyr 2014, for a conceptual model of lion metapopulation management in South Africa). Restoring naturally occurring metapopulations is somewhat easier in that animals do not need to be moved provided that corridors of connectivity are maintained. ...
... This was one of the drivers of the Yellowstone wolf reintroduction (Smith and Bangs 2009), as well as the reintroduction of top-order predators to many sites in South Africa (Hayward et al. 2007b). Indeed, the management of reintroduced predators in South Africa has now moved beyond simply determining when the abundance of a species exceeds the amount of food available to it, to a situation whereby natural dispersal processes can now be mimicked in management activities by translocating surplus animals out of confi ned populations (Ferreira and Hofmeyr 2014). ...
... Big cats, one of the groups of predators that dominate a large part of the world's ecosystems, have managed to evolve different techniques to bring down their prey, which has provided them with their great ecological success [28]. Within this taxonomic group, we strictly include those cats that roar, that is, the four species of the genus Panthera (lion, leopard, jaguar and tiger). ...
... Lions (Panthera leo), cats with a markedly gregarious character, have aroused the interest of scientific groups investigating their strategies and success in hunting, in which females are the main protagonists [28][29][30][31][32][33]. These tend to organize themselves into groups that pursue their prey (medium-sized mammals, mainly ungulates), while males take an active part in hunting actions when they do not belong to any defined group of congeners or in cases where the prey is of considerable size (buffalo, hippopotamus or elephants, for example) as success lies in the coordinated cooperation of several group members [34]. ...
Full-text available
The individuals engaged in predation interactions modify their adaptation strategies to improve their efficiency to reach success in the fight for survival. This success is linked to either capturing prey (predator) or escaping (prey). Based on the graphic material available on digital platforms both of public and private access, this research aimed to evaluate the influence of those animal- and environment-dependent factors affecting the probability of successful escape of prey species in case of attack by big cats. Bayesian predictive analysis was performed to evaluate the outcomes derived from such factor combinations on the probability of successful escape. Predator species, age, status at the end of the hunting act, time lapse between first attention towards potential prey and first physical contact, prey species and the relief of the terrain, significantly conditioned (p < 0.05) escape success. Social cooperation in hunting may be more important in certain settings and for certain prey species than others. The most parsimonious model explained 36.5% of the variability in escaping success. These results can be useful to design translatable selective strategies not only seeking to boost predation abilities of domestic felids for pest control, but also, biological antipredator defence in potential domestic prey of big cats.
... Translocating lions into/out of populations to mimic the natural metapopulation processes of immigration, emigration, births and deaths (Funston, 2008;Slotow and Hunter, 2009;Hayward and Kerley, 2009;Trinkel et al., 2010;Miller et al., 2013;Ferreira and Hofmeyr, 2014;Miller et al., 2015) Genetic Rescue/ Augmentation ...
Full-text available
As the top predator in African ecosystems, lions have lost more than 90% of their historical range, and few countries possess strong evidence for stable populations. Translocations (broadly defined here as the capture and movement of lions for various management purposes) have become an increasingly popular action for this species, but the wide array of lion translocation rationales and subsequent conservation challenges stemming from poorly conceived or unsuitable translocations warrants additional standardized evaluation and guidance. At their best, translocations fill a key role in comprehensive strategies aimed at addressing the threats facing lions and fostering the recovery of wild populations in their historic range. At their worst, translocations can distract from addressing the major threats to wild populations and habitats, divert scarce funding from more valuable conservation actions, exacerbate conflict with humans in recipient sites, disrupt local lion demography, and undermine the genetic integrity of wild lion populations in both source and recipient sites. In the interest of developing best practice guidelines for deciding when and how to conduct lion translocations, we discuss factors to consider when determining whether a translocation is of conservation value, introduce a value assessment for translocations, and provide a decision matrix to assist practitioners in improving the positive and reducing the negative outcomes of lion translocation.
... In South Africa cheetah, Acinonyx jubatus, occur mainly in fenced reserves (Buk et al., 2018). Animals within such fenced reserves often experience elevated competition, as the fences constrain their movements (Ferreira & Hofmeyr, 2014). Alternatively, fenced predator populations may be at higher densities due to fewer external sources of mortality (Packer et al., 2013). ...
en Competition occurs between species for shared resources and the subordinate species can reduce this by avoiding competitors, either proactively or reactively. Cheetah are subordinate members of the African large carnivore guild, vulnerable to losing resources or being killed by larger carnivores. We directly investigate how cheetah space and habitat use are affected by the reintroduction of lion 9 years after cheetah reintroduction. We explored whether cheetah respond on the spatial scale proactively (contracting home range, low overlap and differences in habitat use), reactively (located further than expected at random from lions) or did not respond to lion presence at given monitoring points. We found that there was no difference in cheetah habitat use nor their home range/core area size. Moreover, cheetah home range/core areas overlapped extensively with that of lions. Revealingly, cheetah were located further than expected at random from lions. Our results support a reactive response by cheetah to lion presence. Using this unplanned experiment, we were able to grow our knowledge on the competition occurring between these two large carnivores, by determining that lions did not displace cheetah except on the fine scale. These results can also be of particular use for managers of reserves or metapopulations who are trying to conserve both species sympatrically. Résumé fr Une compétition liée aux ressources partagées se produit entre les espèces, et les espèces subordonnées parviennent à l’atténuer en évitant les concurrents de manière proactive ou réactive. Les guépards font partie d’un groupe subordonné au sein de la guilde des grands carnivores africains, et sont susceptibles de perdre des ressources ou d’être tués par de plus grands carnivores. Notre étude se penche sur la façon dont l’espace et l’utilisation de l’habitat du guépard sont affectés par la réintroduction du lion, qui a eu lieu neuf ans après la réintroduction du guépard. Nous avons tenté de déterminer si les guépards réagissaient à l’échelle spatiale de manière proactive (diminution de l’étendue du domaine vital, faible chevauchement et différences dans l’utilisation de l’habitat), de manière réactive (habitat situé à une distance plus importante que prévu, au hasard de la présence de lions) ou ne réagissaient pas à la présence des lions au sein de sites de surveillance donnés. Nous n’avons constaté aucune différence dans l’utilisation de l’habitat des guépards ou dans la taille de leurs domaines vitaux ou aires principales de répartition. Par ailleurs, les domaines vitaux ou aires principales de répartition des guépards chevauchaient largement ceux des lions. Fait révélateur, des guépards ont été localisés à une distance plus importante que prévu, au hasard de la présence de lions. Nos résultats confirment une réponse réactive des guépards face à la présence des lions. Grâce à cette expérience non planifiée, nous avons été en mesure d’approfondir nos connaissances sur la compétition entre ces deux grands carnivores, en établissant que la présence de lions ne constituait pas un facteur de déplacement des guépards, sauf cas exceptionnels. Ces résultats peuvent s’avérer particulièrement utiles pour les gestionnaires de réserves ou de métapopulations qui tentent de conserver les deux espèces de manière sympatrique.
... Additionally, females that visited the sites without cubs appeared to be in oestrus and looking for mating opportunities. This kind of knowledge can help with population and individual management, as needed in small fenced reserves, providing information that can be otherwise difficult or costly to obtain (Ferreira & Hofmeyr, 2014 ...
Scent‐marking is an important form of communication for solitary species, as the information remains for the receiver after the sender has left the area. Individuals of different demographic or dominance classes (ranks) need to provide different information to conspecifics. Therefore, the use of scent‐marking sites is of particular value for solitary species, but their functional role may vary between demographic classes. Using camera traps, we video‐recorded the behaviour of cheetahs at scent‐marking sites, assessing how individuals of different sex and social rank use the sites for different roles. We tested hypotheses that these sites are for territorial marking, reproductive signalling and information gathering. We found that females visit the sites infrequently, apparently to signal oestrous events, while males visit the sites frequently for territorial marking and information gathering. Among males, dominant individuals scent‐marked the most and reacted to female visitation by vocalizing. In contrast, subordinate males, although also visiting the sites, were not recorded scent‐marking nor vocalizing. These differences in the functional use of scent‐marking sites among sex and social rank support the hypotheses that cheetahs use these scent‐marking sites for both territorial and reproductive signalling, as well as to gather information.
... Cheetahs, Acinonyx jubatus, in South Africa occur mainly in fenced reserves (Buk et al., 2018). Reserves with predator-proof fencing often experience higher levels of competition, as they constrain movements of animals (Ferreira & Hofmeyr, 2014). In addition, tourism, which is often a driving force behind these reserves (Lindsey et al., 2007), favours stocking charismatic animals, such as predators, in higher than 'normal' densities to increase sighting opportunities (Maciejewski & Kerley, 2014). ...
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Competition occurs between species for shared resources. Subordinate species employ resource selection to shift their resource use away from that of dominant species in order to avoid the negative consequences of competition. Only in Africa is the post-Pleistocene large carnivore guild intact, consisting of lions Panthera leo, spotted hyaenas Crocuta crocuta, leopards Panthera pardus, cheetahs Acinonyx jubatus, and African wild dogs Lycaon pictus. Therefore, only in Africa can we explore how large carnivores co-exist with one another in a diverse community. Cheetahs are a subordinate member of the large carnivore guild due to their small stature and solitary nature. However, we still do not fully understand how competition shapes cheetah behaviour and resource use. I used cheetahs as a model subordinate predator to determine the behavioural responses and resource selection of cheetah in response to assumed competition from other large carnivores. I experimentally explored the behavioural response of cheetah to large carnivore scent sources (scats) and their presence at cheetah scent-marking sites to test for avoidance of such cues. Moreover, using unplanned experiments based on the resource use of cheetahs in the absence and then presence of lions (assessing space and time use by cheetah) and African wild dogs (assessing prey use by both species), I evaluated resource selection by cheetahs as a way to reduce competition with these large carnivores. In addition, I measured spatial and temporal partitioning between cheetahs and all four large carnivores at camera trap sites. Finally, I assessed whether cheetahs responded to competitors using a long term proactive response or a short term reactive response. I found that all four of the other large carnivores in the guild shaped cheetahs resource use, however, all did so on different axes in accordance with the niche complementarity hypothesis. Lions and spotted hyenas were avoided through time, African wild dogs through space and prey use, and lions and leopards on the spatiotemporal axis. Moreover, I show that cheetahs utilize a reactive response to competition that allows them to avoid risk while still obtaining necessary resources.
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In an effort to restore parts of their historical geographical range, and in recognition of their ability to restore ecosystem functioning and of the financial benefits they can provide through ecotourism, large carnivores have been reintroduced in many protected areas from which they were previously extirpated. Similar to dispersing animals, translocated individuals often undertake long-distance exploratory movements before establishing home ranges. Post-release monitoring of reintroduced carnivores is common, but the mechanisms of population establishment are rarely examined, limiting our understanding of reintroduction success. We monitored survival and post-release movements of seven cheetahs Acinonyx jubatus reintroduced to Liwonde National Park, Malawi, to evaluate early population establishment. Exploratory phases post-release lasted 29–174 days. Duration of pre-release holding periods in the boma had no significant effect on daily distance moved. Males travelled significantly farther and established home ranges later than females. All cheetahs showed release site fidelity and all females birthed their first litter within 4 months of release. Within 2 years of reintroduction, the newly established population consisted of 14 cheetahs, with demographic attributes similar to those recorded in the source populations. Based on individual settlement, survival and reproduction rates, we deemed this reintroduction successful in re-establishing a breeding population of cheetahs in Liwonde. Our findings suggest the drivers of settlement and population establishment for reintroduced cheetahs are complex, highlighting the importance of assessing and reporting post-release monitoring data.
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Large charismatic predators have been reintroduced to several fenced reserves (<1000 km 2); consequently, their populations can be at risk to declining genetic diversity and catastrophic events such as disease. Welgevonden Game Reserve (WGR) experienced an outbreak of canine distemper virus (CDV) in its lion (Panthera leo) population between October 2015 and February 2016. The decline was attributed to both direct CDV mortalities and indirect management interventions with a population decrease of 93%. In addition, the first documented cases of CDV were recorded in a brown hyaena (Hyaena brunnea) population. This article shares the management of a disease outbreak in WGR's lion population. It discusses the population history, the making of clinical observations leading to diagnoses and recommends strategies that can be implemented to prevent extirpation. The sourcing of new lions from established meta-populations, and the use of vaccinations to proactively prevent future outbreaks in managed wild lions is discussed. In less than two years the population has recovered. This paper provides a case report for wildlife practitioners of small populations of managed wild lions.
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The reproductive biology of a pride of lions (Panthera leo) was studied on the 8500 ha Karongwe Game Reserve from 1999 to 2005. Over this period, the pride consisted of between four and 11 lions with a paired coalition of adult males during the first three years and a single adult male for the next three years. We recorded shorter than normal interbirth intervals, high birth rates of 1.3 cubs/lioness/year, very high cub survival rates, and subadults leaving the pride at young ages. This translated into substantially faster growth rates than are typical in large lion populations in ecologically similar circumstances such as Kruger National Park, but are similar to those of lions in Serengeti National Park. These demographic characteristics were probably induced initially by a lack of intense intraspecific competition and high prey availability, but population stability was maintained through the removal of young subadults by management. Interestingly, variability in conception rates between lionesses resulted in lower growth rates than have been found in other similar reintroduced lion populations. These demographic traits characterize many of the small reintroduced lion populations, and call for appropriate management to avoid the consequences of high predator densities.
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Diverse political, cultural and biological needs epitomise the contrasting demands impacting on the mandate of the South African National Parks (SANParks) to maintain biological diversity. Systems-based approaches and strategic adaptive management (learn by doing) enable SANParks to accommodate these demands. However, such a management strategy creates new information needs, which require an appropriate analytical approach. We use conceptual links between objectives, indicators, mechanisms and modulators to identify key concerns in the context of and related to management objectives. Although our suggested monitoring designs are based mostly on defined or predicted underlying mechanisms of a concern, SANParks requires inventory monitoring to evaluate its key mandate. We therefore propose a predictive inventory approach based on species assemblages related to habitat preferences. Inventories alone may not always adequately serve unpacking of mechanisms: in some cases population size needs to be estimated to meet the information needs of management strategies, but actual population sizes may indirectly affect how the species impact on other values. In addition, ecosystem objectives require multivariate assessments of key communities, which can be used in trend analysis. SANParks therefore needs to know how to detect and define trends efficiently, which, in turn, requires precision of measures of variables. Conservation implications: Current research needs with regard to monitoring should focus on defining designs to yield optimal precision whilst taking methodology, survey trade-offs and analytical approaches into account. Use of these directives and research will guide monitoring during evaluation of SANParks objectives at various scales.
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In South Africa efforts are currently being made to manage several sub-populations of African wild dogs (Lycaon pictus) occurring in isolated, fenced reserves, as a meta-population. This study represents an attempt to estimate the minimum reserve size for the reintroduction of a pack of wild dogs, as a sub-population. Minimum area requirements were based on the area required to support an adequate population of the most important prey species in the diet of a pack of wild dogs. A pack size of five is the threshold below which reproductive failure is likely, and the area requirements of five wild dogs are estimated to be 65 km2 in northern, 72 km2 in eastern and 147 km2 in northeastern South Africa. The presence of perimeter fencing at release sites is a potentially complicating factor, however, as in some cases wild dogs learn to use fences as a hunting tool, permitting the capture of larger prey than is normal. In the event of this happening, larger areas may be required to prevent local population declines in preferred prey species. In general, the use of larger areas is advisable to allow for variation in prey population sizes and the prey profile of wild dogs post-release, and would also be necessary if wild dogs are to be reintroduced into an area with existing populations of lions and spotted hyaenas.
We present our most recent implementation of an individual-based lion population model. The model is built using a structure that includes graphical user interaction and feedback at a fundamental level, to maintain flexibility and ease of use without getting mired in programming details. We describe how object-orientated programming structures may be used to this end. Our structure allows for tight integration between the model and the user interface, including seamless user interaction at any point during the simulation. In addition, the simulation model may be decoupled from the user interface, and large, efficient simulations may be performed. The result is an individual-based model that, despite its complexity, is easy to use, easy to maintain and easy to extend. In fact, it has the potential to be used by a much wider group (including other researchers, managers and conservation advocates) than just the researcher who developed it. We include a discussion of the importance of a robust random number generator in simulation models such as this.