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The Decline in the Lion Population in Africa and Possible Mitigation Measures

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The African lion Panthera leo is under threat. Over the last century the lion has lost about 82 % of its former distribution range, and recent estimates suggest that there are 23,000–38,000 free-ranging lions living in 68 mostly geographically isolated areas. Approximately 24,000 lions are in strongholds, but about 6000 lions are living in populations with a high risk of local extinction, of which about 3000 are in West and Central Africa. Particularly critical is the situation in West Africa, with perhaps only 400 individuals, of which less than 250 are adults. Main threats for lions are habitat loss through agricultural development and human settlement, depletion of prey populations, human–wildlife conflict, epidemics and diseases and trade of lion parts. Conflict mitigation between humans and lions, mainly investigated in rural areas in West and East Africa, have successfully reduced livestock losses and subsequently reduced the number of lions killed. However, mitigation measures alone might fail to secure the critically endangered lion population in West Africa, where translocations and reintroductions might be necessary to counteract genetic impoverishment. Despite encouraging human wildlife coexistence, other approaches for effective long-term conservation of African lions, and the lions’ prey species, favor the separation of land used by humans and conservation areas through the erection of fences. Fences have already been erected in many areas in southern Africa, where lion populations are still viable. Ecotourism and trophy hunting can also be beneficial for lion conservation. However, revenues and user rights over wildlife have to be devolved to local people. This has been successfully implemented in Namibia, which is one of the few countries where lion populations are currently increasing. Although trophy hunting can be a useful conservation tool for conserving lions, it can also be a threat. Therefore, improved regulations and management of lion hunting to prevent negative impacts on hunted lion populations is necessary in many countries. If the downwards trend in lion numbers continues, it is questionable if there will be still lions around in the next century.
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45© Springer International Publishing Switzerland 2016
F.M. Angelici (ed.), Problematic Wildlife, DOI 10.1007/978-3-319-22246-2_3
Chapter 3
The Decline in the Lion Population in Africa
and Possible Mitigation Measures
Martina Trinkel and Francesco Maria Angelici
Introduction
During the late Pleistocene lions Panthera leo spp. ranged all over Africa, Eurasia,
and some parts of North America (Steele 2007 ; Barnett et al. 2009a , b ). Today
African lions only occur in Sub-Saharan Africa, where they have suffered a marked
reduction of range and population sizes (Nowell and Jackson 1996 ). Over the last
century the lion, classifi ed as vulnerable in the IUCN Red List (Bauer et al. 2012 ),
has lost about 82 % of its former distribution range (Ray et al. 2005 ). There are
strong indications that lion populations are declining: current population estimates
range from 23,000 to 38,000 free-ranging lions living in 68 lion areas (Fig. 3.1 )
(Chardonnet 2002 ; Bauer and Van Der Merwe 2004 ; Riggio et al. 2013 ).
Approximately 24,000 lions are in strongholds, but about 6000 lions are living in
populations with a high risk of local extinction, of which about 3000 are in West and
Central Africa (Riggio et al. 2013 ).
Recently, Bauer et al. ( 2015 ) assessed the trend of 47 relatively well-monitored
lions in Africa, and found an alarming population decline of about 38 % over 21 years
(1993–2014). There were signifi cant regional differences in lion population trends: in
southern Africa, lion populations grew by 22 %. In contrast, lion populations decreased
by 57 % in East Africa, and 66 % in West and Central Africa (Bauer et al. 2015 ).
In general, the largest lion populations occur in East and southern Africa with
about 11,000 and 10,000 individuals, respectively (Bauer and Van Der Merwe
2004 ). Many of the large lion populations in southern Africa have been stable
M. Trinkel (*)
School of Life Sciences, University of KwaZulu-Natal , Westville , Durban 4041 , South Africa
e-mail: martina.trinkel@gmail.com
F. M. Angelici
Fondazione Italiana per la Zoologia dei Vertebrati (FIZV) , Via Cleonia 30, Scala C , Rome
00152 , Italy
e-mail: francescomariaangelici@gmail.com
46
(or even increasing) over the last decades, where conservation of lions has benefi ted
from the development of the ecotourism and trophy hunting industry (Chardonnet
2002 ). In West and Central Africa the most severe decline of the lions’ range has
occurred with estimates of only 850–950 individuals in 2001/2002 (Bauer and Van
Der Merwe
2004 ). More recently, the West African lion population has declined
drastically in terms of both numbers and range, resulting in a few small, geographi-
cally distant remaining populations (Henschel et al. 2014 ). In most non-protected
areas in West and Central Africa, however, the lion has disappeared (Bauer and Van
Der Merwe 2004 ).
All over Africa, the dramatic decline of the lion is mainly due to an increasing
confl ict between wildlife and humans over space (Nowell and Jackson 1996 ).
Agriculture, human settlement, depletion of prey populations, and direct persecution
Fig. 3.1 Historic and present distribution of the lion Panthera leo in Africa
M. Trinkel and F.M. Angelici
47
have led to shrinking habitat for large carnivores (Nowell and Jackson 1996 ). Lions,
therefore, more and more live in patchy and geographically isolated conservation
areas (Nowell and Jackson 1996 ; Chardonnet 2002 ). Even national parks and pro-
tected areas cannot fully protect lions due to bushmeat hunting (Lindsey et al. 2013 )
and severe confl icts with humans on reserve borders (Woodroffe and Ginsberg
1998 ). When communities do not receive revenue from the species, it is diffi cult to
nd solutions for human–wildlife confl icts (Jorge et al. 2013 ). Therefore, it is vital
for lion conservation to offer incentives for local communities and individual land-
owners, so that conserving lions becomes economically sustainable and profi table
(Nelson et al. 2013 ).
In this review, we address problems that threaten the African lion population
such as loss of habitat and connectivity, prey depletion, human–wildlife confl ict,
epidemic and diseases and the trades of lion parts. We summarize possible solutions
to mitigate human–wildlife confl ict, which were mainly investigates in parts of
East, West, and Central Africa. Further, we address the issue of fencing conserva-
tion areas to separate humans and wildlife and the development of ecotourism and
trophy hunting . In West Africa, where the species is Critically Endangered, we
highlight which measures have to be taken quickly to prevent localized extinction.
Taxonomy
Analysis of mitochondrial DNA of lions from West and Central Africa showed that
these populations are more closely related to Asiatic lions rather than to lions in East
and southern Africa (Bertola et al. 2011 ). This may have resulted from an extinction of
lions during the Pleistocene, followed by a re-colonization of North Africa, just after
the late Pleistocene (Bertola et al. 2011 ). Re-colonization might have continued towards
West and Central Africa (i.e., from Yankari Game Reserve, Eastern Nigeria to South
Sudan) derived from the Middle East and Asia (Bertola et al. 2011 ). Besides lion popu-
lations in West and Central Africa, some other African lion populations, e.g., lions in
Ethiopia , differ morphologically and genetically from lions in East and southern Africa
(Bruche et al. 2013 ). For these reasons, taxonomy and sub-specifi c nomenclature of
African lions should be reorganized, which would ultimately affect conservation strat-
egies of the species both in situ and ex situ (Bertola et al. 2011 ; Henschel et al. 2014 ).
Threats
Habitat Loss, Isolation, and Prey Depletion
Large carnivores have suffered a marked decline due to increased human population
pressure leading to habitat destruction (Patterson
2004 ), and patchy and geographi-
cally isolated conservation areas (Nowell and Jackson 1996 ). Human population
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
48
growth and the expansion of agriculture has caused massive declines of almost all
larger mammalian carnivores, resulting in range collapses down to a few per cent of
their original distribution (Woodroffe et al. 2005 ).
In recent decades, lion distribution was drastically reduced and fragmented
(Riggio et al. 2013 ). Nowadays, many areas where lions still occur are isolated
without any connection to each other, in particular in West Africa and Ethiopia
(Fig. 3.1 ). Many populations are small and may have insuffi cient genetic variability,
which is essential for maintaining a healthy population in the long term (Björklund
2003 ). The most important protected areas for the conservation of large mammals,
including the African lion, are concentrated in southern and East Africa (Wegmann
et al. 2014 ). In West Africa, where lion numbers are low and ranges are isolated,
conservation actions are urgently needed (Henschel et al. 2014 ). Based on recent
surveys in West Africa, from Senegal to Western Nigeria (Tende et al. 2014 ), there
are about 400 lions in the wild representing less than 250 mature individuals. These
lions are concentrated in only four protected areas, forming two separate blocks
(Fig. 3.1 ). To counteract genetic impoverishment of the critically endangered lion
population in West Africa, translocations and reintroductions might be necessary
(Trinkel et al. 2008 ). However, it is likely that there are more small lion populations
in areas which have not yet been investigated and/or are currently monitored, i.e., in
Guinea, Ivory Coast, and Ghana (Henschel et al. 2014 ; Angelici et al. submitted).
Case Study: Lions in the Mole National Park (Ghana)
The Mole National Park ( MNP , size: 4600 km 2 ) in Ghana is one of the most impor-
tant protected areas in West Africa, where lions have always been present (Angelici
and Petrozzi 2010 ; Angelici et al. 2015 ). Recently, it was considered that lions in
MNP and possibly in whole Ghana are extinct (Burton et al. 2011 ; Henschel et al.
2014 ). In 2004, a male lion was shot in MNP and this individual was thought to be
the last lion of the area. In order to evaluate if there are remaining lions in the MNP
and surrounding areas, the project “The Pride of Ghana” was initiated in 2005
(Angelici and Petrozzi 2010 ; Angelici et al. 2015 ). It is a cooperation between the
Forestry Commission of the Ghanaian Ministry of Lands and Forestry, the Italian
NGO “Ricerca e Cooperazione,” and the University of Tuscia, Viterbo, Italy.
The methods used in this project are camera traps, sound playbacks, and interviews
with local people. Since 2005, lion tracks, prey presumably killed by lions, and
roars were regularly registered by park staff and researchers (Angelici et al. 2015 ).
Due to the large size of the MNP and the extremely scarce population, lions are dif-
cult to count in this area (Myers 1975 ). Therefore, close collaboration with park
staff and local people is of tremendous importance. Such collaborations with local
people are often underrepresented or even ignored in fi eld research resulting in the
loss of important information (Black et al. 2013 ).
Wegmann et al. ( 2014 ) argued that for conservation of large mammals, i.e.,
ungulates and large carnivores, it is essential that protected areas throughout Africa
are connected with each other. Even when connected, each protected area is vulnerable
M. Trinkel and F.M. Angelici
49
to habitat change which could limit the effectiveness of connectivity in a network of
protected areas, and thus increase the risk of the extinction of species (Wegmann
et al. 2014 ). Important parameters for connectivity are size and location of protected
areas and, in general, habitats are better preserved inside rather than outside pro-
tected areas, where fast deterioration of habitat often occurs (Wegmann et al. 2014 ).
Thus, it is essential to not only preserve protected areas, but also non- protected
areas (Wegmann et al. 2014 ). To effectively conserve lions, it would be of high
value to connect protected areas, whilst at the same time considering genetic differ-
ences (Bertola et al. 2011 ). In West and Central Africa, and in Ethiopia, where
remaining lion populations are extremely small and isolated (Fig. 3.1 ), protected
areas should be increased and connected through ecological corridors to maintain
genetically viable populations. Failing this (or while it is being implemented), trans-
location schemes may be needed in the short term to conserve or increase genetic
diversity (Hunter et al. 2007 ; Trinkel et al. 2008 ). A recent study of lions in Hwange
National Park, Zimbabwe, demonstrated the importance of smaller-scale connectiv-
ity, showing that the genetic pattern of the Hwange lion population can be attributed
to still existing high levels of habitat connectivity between protected areas (Morandin
et al. 2014 ). In areas where direct connectivity cannot be employed, genetic con-
nectivity could be achieved with meta-population management (Akçakaya et al.
2006 ; Gusset et al. 2008 ; Slotow and Hunter 2009 ; Lindsey et al. 2011 ).
Lions depend on medium and large prey species (Van Orsdorl et al. 1985;
Hayward and Kerley 2009 ). Large mammals, however, were found to decline dra-
matically in most African countries (Craigie et al. 2010). Between 1970 and 2005,
wildlife population abundance severely declined in West Africa (85 %) and East
Africa (52 %); only southern Africa was able to maintain their wildlife populations
(Craigie et al. 2010). In some areas, bushmeat trade, the illegal hunting and com-
mercialization of bush meat, is a more pressing problem than the loss of habitat
(Wilkie et al. 2011 ). Due to disappearing wildlife in unprotected areas (Newmark
2008), illegal bushmeat hunting has been increasing dramatically in protected
savannah areas (Lindsey et al. 2013 ). The hunting and trade of bushmeat leads to
edge effects around protected areas, disproportionate declines of particular spe-
cies, and severe population declines in areas with inadequate anti-poaching
(Lindsey et al. 2013 ), which all negatively affects lion population size (Packer
et al. 2014 draft).
Human–Lion Confl ict
When wildlife has little or no value outside protected areas, it rapidly disappears
through habitat loss and direct persecution (Prins and Grootenhuis 2000 ).
Anthropization, including expansion of agriculture, has caused massive declines of
almost all larger mammalian carnivores (Woodroffe et al. 2005 ). In West and Central
Africa, lion population density is low, with approximately 1–3 lions/100 km
2 (Bauer
and Van Der Merwe 2004 ). Such low population densities are due to low prey
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
50
densities (Hayward et al. 2007a ; Bauer et al. 2008 ) and high human densities,
which results in frequent and intense interactions between humans and lions (Bauer
et al. 2010 ). Lions frequently kill livestock and are subsequently trapped, shot, or
poisoned (Bauer and De Iongh 2005 ).
Even in conservation areas, large carnivores are not suffi ciently protected.
Poaching is a problem (e.g., Weladji and Tchamba 2003 ; Hilborn et al. 2006 ) and
confl ict with people at reserve borders negatively infl uences populations over wide
areas (Weladji and Tchamba 2003 ). Increasing security through elevated anti-
poaching can effectively reduce bushmeat hunting inside protected areas (Lindsey
et al. 2013 ), which in turn reduces the decline of lions and wildlife populations in
general. Edge-related mortality at reserve boundaries can lead to population declines
or even extinction (Scheepers and Venzke 1995 ; Woodroffe and Ginsberg 1998 ;
Balme et al. 2010 ). Problem animal control is a major source of mortality for lions
outside of protected areas (Woodroffe and Ginsberg 1998 ), and is often carried out
as a response to lions that prey on livestock (Kötting 2002 ; Woodroffe et al. 2006 ).
For example, in the Kgalagadi Transfrontier Park, South Africa/Botswana, Van
Vuuren et al. ( 2005 ) argued that, due to problem animal control, the lion population
probably cannot be sustained in the long term.
In Namibia, the largest lion population occurs in the Etosha National Park, an
arid habitat of immense size (approximately 23,000 km
2 , about the same size as the
Kruger National Park, South Africa) with an estimated population size of about 320
individuals (Stander 1991 ). Although lions are protected within the park, confl ict is
a problem where Etosha borders on surrounding commercial and communal farm-
land. More than 1000 lions were shot or destroyed in the last 20 years (Etosha
Ecological Institute). These numbers, however, include only reported cases, and the
actual number of killed lions is thought to be higher (Tana Burger, Personal
Communication). Kötting ( 2002 ) investigated livestock losses due to problem ani-
mals, mainly lions and spotted hyenas, on commercial farms along the southern
boundary of the Etosha National Park. Farmers on average lost livestock worth
2000€ per year to lions, and on average, one lion was shot on each farm per year.
There existed a linear relationship between livestock losses and the number of
destroyed lions, with one outlier: one farmer destroyed 400 lions in 55 years and
this particular farmer is known to lure lions out of the park to subsequently destroy
them (Kötting 2002 ). Despite such massive losses of lions through border confl icts,
Trinkel ( 2013 ) found that the lion population size inside Etosha remained stable
over a 20-year study. However, population size alone is not a good measure for the
health of a population, especially not for lions, who have the ability to quickly
recover from disturbances (Smuts 1978 ; Packer et al. 1990 ; Trinkel 2013 ). Recently,
Trinkel et al. (submitted) found that the persecution of sub-adult males on farmland
bordering Etosha was disproportionately high, and although it did not affect lion
population size, it affected social dynamics of lions inside the national park. Due to
a lack of competition from sub-adult males, Etosha males hold tenure over prides
for nearly 9 years, which is four to fi ve times longer than pride males in other
ecosystems (Packer et al.
1988 ; Yamazaki 1996 ; Bauer et al. 2012 ; Trinkel et al.
submitted). As young females usually fi rst giving birth at 3–4 years of age and give
M. Trinkel and F.M. Angelici
51
birth every 2 years (Packer et al. 1988 ), there is consequently a high risk of them
mating with direct relatives (Trinkel et al. 2010 , submitted). Trinkel et al. (submitted)
argue that if managed properly (Packer et al. 2009 ; Balme et al. 2012 ), wildlife
tourism and sustainable utilization through trophy hunting in these areas may benefi t
species conservation (Selier et al. 2014 ) and could thus contribute to the genetic
health and sustainability of the Etosha lion population.
Epidemics and Animal Diseases
Besides direct persecution, close contact between livestock and wildlife bears other
risks: infectious diseases can have a dramatic impact on wildlife populations and is
important in carnivore conservation (Scott 1988 ). Viruses have caused major
declines in populations of large carnivores (Young 1994 ), such as canine distemper
(Roelke-Parker et al. 1996 ), rabies (Sillero-Zubiri et al. 1996 ), and bovine tubercu-
losis (Keet et al. 1996 ). However, Ferreira and Funston ( 2010 ) evaluated the effect
of bovine tuberculosis on estimates of lion density and survival in the Kruger
National Park, South Africa, and found that the size of the lion population was sta-
ble, despite the high prevalence of bovine tuberculosis. Detailed analysis of the
canine distemper virus epidemic in lions of the Serengeti and the Ngorongoro
Crater, Tanzania, revealed that canine distemper is only a problem when lions are
coinfected with high levels of Babesia (Munson et al. 2008 ).
Diseases can be transmitted through domestic animals, such as dogs or livestock,
and can be lethal for populations of wild species such as lions (e.g., Roelke-Parker
et al. 1996 ; Packer et al. 1999 ). Canine distemper—probably transmitted by dogs
via wild canids and subsequently onto lions—killed over 35 % of lion in the
Serengeti, East Africa, in 1994 (Roelke-Parker et al. 1996 ; Carpenter et al. 1998 ),
but only if the lions were coinfected with Babesia (Munson et al. 2008 ). Bovine
tuberculosis is a pathogen of growing concern in free-ranging wildlife in southern
Africa (Trinkel et al. 2011 ) and was shown to be transmitted from livestock to wild-
life populations (Michel et al. 2006 ). For example, in the Kruger National Park,
South Africa, bovine tuberculosis has resulted in the mortality of buffalo Syncerus
caffer , lion, and cheetah Acinonyx jubatus (Keet et al. 1996 ; De Lisle et al. 2002 ;
Ferreira and Funston 2010 ). To prevent such fatal consequences, it is important that
livestock and pets of humans living close to wildlife be vaccinated against diseases
that can threaten wildlife populations (Harder et al. 1995 ). However, controlling
canine distemper with a large-scale dog vaccination program around the Serengeti,
Tanzania, did not prevent the spread of the disease onto lions (Viana et al. 2015 ).
Despite the fact that direct vaccination of threatened wildlife has been heavily
debated (Cleaveland et al. 2006 ), there has been considerable progress in the devel-
opment of safe vaccines for large carnivores (Pardo et al. 1997 ; Wimsatt et al. 2003 ).
Furthermore, mathematical models suggest that vaccinating only about 30–40 % of
the individuals in a large carnivore population would be suffi cient to effectively
protect the population from serious diseases (Haydon et al. 2006 ; Vial et al. 2006 ).
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
52
Therefore, instead of a large-scale vaccination program for domestic animals, it
might be more effective to directly vaccinate the lions themselves (Viana et al.
2015 ). In small reserves in South Africa, lions are frequently translocated to restore
and/or maintain their health status and genetic diversity (Trinkel et al. 2008 , 2011 ).
To prevent negative consequences, lions are carefully screened for diseases prior to
translocations (Trinkel et al. 2011 ).
Trade in Live Pets, Bones, Meat, and Skins
A problem that could have major implication on lion populations in the near
future is an increasing trade of lion bones, skins, dried meat, and other parts of the
lion’s body for use in traditional medicine, mostly in Asia (Ellis 2005 ). As the
tiger Panthera tigris is becoming rarer, the lion is replacing the tiger as an ingredi-
ent for traditional medicine and magic (Ellis 2005 ; Morell 2007 ; Gratwicke et al.
2008 ). Many captive lion breeders in South Africa obtain high incomes from the
sale of lion carcasses to China, Vietnam, and other Asian countries (Lindsey et al.
2012a ; Nowell 2012 ). Such trade, however, is dangerous, because it is diffi cult to
control which lions were killed legally and which ones were poached. Poaching
certainly is widely present in both the offi cial and the illegal markets (Ellis 2005 ;
Zhang et al. 2008 ). If the trade is not carefully controlled, lions could decline as
dramatically as the tiger (Dinerstein et al. 2007 ). Therefore, any trade has to be
seriously controlled and severe sanctions for poachers have to be installed
(Gratwicke et al. 2008 ).
The trade of lions for pets seems to be declining (Hemley 1994 ). This is mainly
due to legalities prohibiting the trade in wild animals for the pet trade (e.g., Nyhus
et al. 2009 ). People who want to keep large carnivores at their home or in cages can
mostly get lions from captive breeders or circuses (Hemley 1994 ; Bush et al. 2013 ).
While this trade is of concern, it has less impact on the wild lion population than the
trade in lion parts.
Possible Mitigations
Human–Lion Confl ict : Possible Solutions
A reduction in livestock losses would ultimately reduce the number of lions and other
predators that are killed benefi tting both people and wildlife (Ogada et al. 2003 ;
Woodroffe et al. 2006 ). Some conservationists, therefore, have investigated the
effects of livestock husbandry for mitigating human–wildlife confl icts along reserve
borders and outside protected areas (Ogada et al. 2003 ; Kissui 2008 ; Jorge et al.
2013 ). For example, Woodroffe et al. ( 2006 ) conducted a study in a non- protected
area on communal land in Kenya, where attentive livestock husbandry is common:
97 % of livestock herds were accompanied by herders (Woodroffe et al.
2006 ).
M. Trinkel and F.M. Angelici
53
The reason for continuing traditional husbandry in this area was mainly the high
risk of livestock being stolen rather than being killed by predators. In such areas,
small changes in livestock husbandry were shown to effectively reduce livestock
losses (Ogada et al. 2003 ; Woodroffe et al. 2006 ). These changes included con-
structing denser boma walls and the presence of domestic dogs at bomas and with
grazing herds (Ogada et al. 2003 ; Woodroffe et al. 2006 ). Lichtenfels et al. (2015)
evaluated fortifi ed bomas just east of Tarangire National Park on the Maasai Steppe,
Tanzania, and found that carnivore attack rates at fortifi ed bomas called Living
Walls were signifi cantly reduced. Living Walls are predator-proof enclosures that
consist of fast-growing, thorny trees as fence posts with chain link fencing
(Lichtenfels et al. 2015). A so-called Lion Guardians program has been successfully
incorporated in the Massai society in Kenya, where lions killing livestock in pasto-
ralist areas are traditionally poisoned and speared, which—besides retaliation—
reinforces the role of warriors (Hazzah et al. 2014 ). Warriors in the Maasai society
are well-respected people, and are responsible for protecting their community and
livestock from carnivore attacks (Hazzah et al. 2014 ). Within the “Lion Guardians”
program, warriors defend their community by “hunting” lions to radio-collar and
monitor them, while receiving economic incentives and cultural recognition (Hazzah
et al. 2014 ). As a result, no lions have been speared in areas where the Lion
Guardians are working since the program started (Hazzah et al. 2014 ). Another
study conducted in Kenya showed that a land use system based on temporary settle-
ments and grazing areas allowed lions to co-occur with people and livestock at
high density (Schuette et al. 2013 ). Dickman et al. ( 2011 ) reviewed the success of
providing economic incentives for local people to tolerate the costs of living with
wildlife. In particular, Dickman et al. ( 2011 ) investigated three major approaches:
(a) the concept of compensation payments, which involves a payment to the live-
stock owner if the suspected incident of livestock depredation is attributed to a
predator; (b) revenue sharing where local people directly benefi t from wildlife as
revenue generated from wildlife tourism are shared among communities; and (c)
conservation payments, which are directly linked, e.g., to the maintenance of preda-
tors on private or community land. Dickman et al. ( 2011 ) found that the best way of
“payments to encourage coexistence” might be a combination of compensation pay-
ments, revenue sharing, and conservation payments. Payments to encourage people
to coexist with wildlife can also be a useful tool to decrease bushmeat hunting
(Lindsey et al. 2013 ).
In West and Central Africa several methods for mitigating confl ict between lions
and humans were investigated, with encouraging results (Bauer et al. 2010 ). For
example, in Pendjari National Park (Benin), enclosures of clay instead of thorny
branches drastically reduced attacks of lions on livestock (Bauer et al. 2010 ). In
Chad, protecting livestock herds with dogs and herdsmen, and creating settlements
with houses surrounding livestock mitigated human–lion confl icts. The use of
magic was found to be intensively practiced throughout West Africa, where rural
people invest considerable amounts of money in the magical protection of their
livestock (Bauer et al.
2010 ). Bauer et al. ( 2010 ) argue that although the effective-
ness of such measures is not relevant, such practices should receive more attention
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
54
when starting discussions with, and fi nding solution for, rural communities. Such
mitigation measures could be used as a sensitisation instrument, e.g., by using
Sourats on community radio (Bauer et al. 2010 ).
Improving livestock husbandry was shown to be a useful tool to mitigate human–
wildlife confl ict in many areas (Ogada et al. 2003 ; Woodroffe et al. 2006 ; Bauer
et al. 2010 ; Lichtenfels et al. 2015; Hazzah et al. 2014 ). However, the effectiveness
of the various techniques depends on the social and cultural susceptibility to such
mitigation measures, and the applicability of each practice has to be evaluated from
case to case. In many parts of Africa traditional husbandry has been abandoned, and
it is therefore questionable if such mitigation measures will be useful in such areas
(Breitenmoser et al. 2005 ) .
Fences for Lion Conservation
Conservation fences are designed to separate biodiversity from factors threatening
it (Hayward and Kerley 2009 ). For effective long-term conservation of African
lions, Packer et al. ( 2013 ) recently recommended the erection of fences to separate
protected areas from land used by humans. Packer et al. ( 2013 ) compared African
lion population densities and population trends in fenced and unfenced reserves
across 42 sites in 11 countries, and found that lion populations in fenced reserves
were signifi cantly closer to their carrying capacity than unfenced populations.
Furthermore, lions in fenced reserves were mainly limited by density dependence
(Packer et al. 2013 ). In contrast, lions in unfenced reserves were highly sensitive to
human population densities in surrounding communities, and these lion populations
were subjected to density-independent factors (Packer et al. 2013 ). Packer et al.
( 2013 ) predicted that nearly half the unfenced populations may decline to near
extinction over the next 20–40 years.
Fences, however, can inhibit or prevent transmigration, i.e., immigration and
emigration , and therefore, fenced reserves are vulnerable to problems associated
with isolated populations (Lindsey et al. 2012b ). Such populations are more suscep-
tible to environmental, demographic, and genetic stochasticity (Caughley 1994 ;
Lindsey et al. 2012b ). Fences can lead to loss of dispersal and migration routes
(De la Bat 1982 ; Creel et al. 2013 ), and prohibit the use of patches of primary pro-
ductivity to wildlife populations (Williamson and Mbano 1988 ; Lindsey et al.
2012b ). For example, the fence around the Etosha National Park reduced the number
of blue wildebeest Connochaetes taurinus from more than 25,000 to 2500 animals
(De la Bat 1982 ). The veterinary fences of Botswana reduced access of zebra Equus
burchelli , blue wildebeest, and other wildlife to water and dry-season grazing areas
and consequently caused a massive number of mortalities (Williamson and Mbano
1988 ; Lindsey et al. 2012b ; Gadd 2012 ). For lion populations, depending on the size
of the enclosed population, fencing often results in the need for genetic and demo-
graphic management of smaller populations via translocations (Trinkel et al.
2008 ;
Miller et al. 2013 ).
M. Trinkel and F.M. Angelici
55
Fencing is becoming an increasingly important conservation tool in southern
Africa to contain animals inside protected areas and limit human–wildlife confl ict
at reserve borders, especially when reserves border on highly populated agricultural
areas (Lindsey et al. 2012b ). In South Africa, for example, all small wildlife reserves
and even the 20,000 km
2 Kruger National Park are fenced by law (Anonymous
2003 ). This fencing has successfully reduced human–lion confl ict (Hayward and
Kerley 2009 ), and can also decrease the impact of bushmeat hunting onto wildlife
populations (Lindsey et al. 2013 ) However, for small areas, the breakdown of
natural processes that regulate lion population growth must also be considered
(Miller et al. 2013 ). Namibia erected a game-proof fence around the 23,000 km
2
Etosha National Park, starting in 1961 and fi nishing in 1973 (Berry 1997 ). However,
this fence has proved inadequate for large predators such lions and spotted hyenas
Crocuta crocuta (Berry 1997 ). Warthogs Phacochoerus africanus and porcupines
Hystrix africaeaustralis dig holes under the fence, which are then frequently used
by lions and other predators to bypass the fence (T. Burger, Personal Communication).
Besides these holes, the maintenance of the 850 km fence is a major problem
(S. Kötting, Personal Communication). As a result, the confl ict between humans
and predators has not been limited (Trinkel et al. submitted), resulting in the killing
of more than 1000 lions in 20 years (Etosha Ecological Institute).
So far, fencing has only been widely employed in a few countries in southern
Africa, mainly because of aesthetic concerns, fi nancial costs, and the impracticality
of enclosing large-scale migratory ungulate populations (Packer et al. 2013 ). In West,
Central, and most of East Africa, the use of fencing in wildlife management is rare
(Lindsey et al. 2012b ). For fencing to be an effective tool for limiting human–wild-
life confl ict, fences have to be well constructed (e.g., electrifi ed) and maintained
(Lindsey et al. 2012a , b ). Fencing, however, has been considered on a case-by-case
basis (Hayward and Kerley 2009 ).
Ecotourism and Trophy Hunting
Finding solutions for human wildlife confl icts is diffi cult, especially when commu-
nities do not receive revenue from the species (Jorge et al. 2013 ; Lindsey et al.
2013 ). Therefore, ecotourism and trophy hunting can be most benefi cial to lion
conservation where revenues and user rights over wildlife are devolved to local
people (Nelson et al. 2013 ). The greatest threat to the sustainability of ecotourism
and trophy hunting on communal land is the failure of governments, tour and hunt-
ing operators to devolve adequate benefi ts to local communities , which reduces
incentives for rural people to conserve wildlife (Lewis and Alpert 1997 ; Mayaka
et al. 2004 ; Child 2005 ; Lindsey et al. 2006 ) and promotes bushmeat hunting
(Lindsey et al. 2013 ).
While West, Central, and East Africa have all witnessed a long-term decline in
large mammal populations, southern Africa has generally experienced substantial
recoveries in wildlife populations across large areas of private land and in some
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
56
cases communal lands (Bond et al. 2004 ; Cragie et al. 2010 ; Nelson et al. 2013 ).
In southern Africa, in particular South Africa, Zimbabwe, and Namibia, revenues
from ecotourism and trophy hunting were largely responsible for the development
of the game-farming industry (Bond et al. 2004 ).
Small Reserves in South Africa
In recent years there has been a rapid increase in the number of small (<1000 km
2 ),
privately owned wildlife reserves in South Africa, many of which have been estab-
lished for ecotourism and for biodiversity conservation (Hayward et al. 2007b ). The
development of the ecotourism industry led to an increase in the reintroduction of
lions. More than 700 wild lions have been reintroduced into 45 small reserves in
South Africa, adding considerable to two naturally occurring lion populations in the
Kruger National Park and the Kgalagadi Transfrontier Park (Miller et al. 2013 ).
However, many of these isolated lion populations have problems with inbreeding
and overpopulation (Grubbich 2001 ; Vartan 2002 ; Trinkel et al. 2008 , 2010 ; Miller
and Funston 2014 ). Rapid population growth of lions is due to high levels of prey
abundance (Vartan 2002 ) and a breakdown of natural processes to regulate lion
population growth (Miller and Funston 2014 ). Due to a lack of guidelines on how to
effectively manage these populations, these populations may be of limited value for
the conservation of this species (Frankham 2009 ; Slotow and Hunter 2009 ).
Therefore, Slotow and Hunter ( 2009 ) suggested to consider a meta-population
management plan for lions in small reserves to insure high longer-term conserva-
tion potential. For wild dogs, meta-population management has already been suc-
cessfully applied for the reintroduction and management of wild dogs in small
reserves in South Africa (Akçakaya et al. 2006 ; Gusset et al. 2008 ). A viable wild
dog population was established consisting of several sub-populations in a network
of small fenced reserves (Davies-Mostert et al. 2009 ; Lindsey et al. 2012b ). Cheetahs
are also managed as a meta-population in South Africa (Lindsey et al. 2011 ). In
2010, a Lion Management Forum was formed to improve the conservation status of
lions in small reserves by implementing a meta-population management plan for
lions in South Africa (Miller et al. 2013 ).
Genetic Isolation and Inbreeding: Translocation as a Conservation Tool
A Case Study from the Hluhluwe iMfolozi Park
Small reserves may have major problems with inbreeding of lions (Vartan 2002 ).
Through translocation, it was shown that it is possible to successfully restore the
genetic health of a small, isolated lion population (Trinkel et al. 2008 ; Miller et al.
2013 ). The native Hluhluwe-iMfolozi Park (HiP) lion population descends from
only three individuals, which repopulated the area in the 1960s (Trinkel et al. 2008 ).
M. Trinkel and F.M. Angelici
57
In 1987 and 1999, the population consisted of about 140 and 80 individuals,
respectively (Maddock et al. 1996 ; Balfour et al., unpublished data). The population
declined since the early 1990 and showed little genetic variation and cub mortality
was found to be high (Stein 1999 ). General poor condition of HiP lions and reduced
immune competence were thought to be associated with inbreeding (Stein 1999 ).
To restore the genetic variation of the inbred HiP lion population, new (outbred)
lions were translocated into the existing population (Trinkel et al. 2008 ): between
1999 and 2001, 16 new lions were translocated into HiP. Three different types of
translocation in four separate releases were undertaken. First, a pride along with its
resident coalition was translocated. Second, a pride of females was translocated.
Third, new females were attempted to bond into preexisting native HiP prides. The
translocations into the existing lion population in the Hluhluwe-iMfolozi Park were
successful (Trinkel et al. 2008 ). They were designed so as to encourage the females
to form four separate prides, but instead they split into six prides comprising of
related and/or unrelated lionesses. Although it appeared as if a cohesive social bond
had developed between all translocated female lions while they were still in the
boma, many individuals separated after release. All translocated females established
stable pride ranges. The three translocated males split into a pair (brothers) and a
singleton, and associated with both native and new females. The pair was more suc-
cessful than the solitary, gaining residence in a pride more easily and maintaining
residence for longer. The native HiP population consisted of about 84 lions in 2000
but crashed to only 20 native individuals and their offspring by 2004, corresponding
to 32 % of the total population. Offspring of translocated and native HiP lions
totalled 29 individuals by the end of 2004 (47 %), and the translocated lions and
their offspring totalled 13 individuals (21 %) (Trinkel et al. 2008 ). Thus, by the end
of 2004, the native HiP population consisted of 20 lions, while translocated lion and
offspring involving with at least one translocated animal comprised 42 animals.
Descendants with at least one introduced lion replaced the entire purebred native
lions by 2006 (Trinkel et al. 2008 ).
The translocations into the existing lion population in the Hluhluwe-iMfolozi
Park were very successful. Such translocations may become an important adaptive
management tool as lion populations become increasingly fragmented (Trinkel
et al. 2008 ).
Commercial and Community Conservancies in Namibia
In Namibia, there has been an ongoing process of transforming former livestock
farmland into wildlife and trophy hunting farms, where both humans and wildlife
benefi t (Göttert and Zeller 2008 ). User rights over wildlife were devolved to private
landowners in the 1960s and 1970s (Bond et al. 2004 ). Since then, large areas of
privately owned commercial cattle farms have been converted into wildlife farms.
Due to such land conservation, wildlife has recovered tremendously and nowadays,
10–20 times more wild ungulates occur on private land than in protected areas
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
58
(Lindsey et al. 2013 ). Starting in the 1990s, Namibia—as the only country in
Africa—devolved full user rights over wildlife to local communities. Such com-
munities are able to establish “communal conservancies” where local residents can
legally hunt wildlife for their own consumption or sell a quota to a hunting operator,
keeping 100 % of the revenue (Jones and Weaver 2009 ; Nelson et al. 2013 ).
Devolving user rights over wildlife to local communities has a large potential to
reduce bushmeat poaching and increase wildlife populations (Lindsey et al. 2013 ).
According to the Conservancy Association of Namibia, “A Conservancy is a legally
protected area of a group of land-occupiers practicing cooperative management
based on a sustainable utilization strategy, promotion of the conservation of natural
resources and wildlife, and the desire to reinstate the original biodiversity with the
basic goal of sharing resources amongst all members” (Shaw and Marker 2010 ).
To date, more than 70 community conservancies covering 160,000 km
2 have
developed, resulting in rapid increases in wildlife populations (Lindsey et al. 2013 ).
Through tourism and hunting, the Kunene lion population in north-western Namibia
has been increasing continuously from about 15 individuals in 1999 to more than
130 individuals in 2010 (Stander 2010 ). Nowadays, the Kunene lion population is
connected with the lion population of Etosha National Park, which holds the largest
lion population in the country (Stander 2010 ; Trinkel et al. submitted). However, for
many commercial wildlife farmers—similar to livestock farmers—large carnivores
are competitors and are heavily persecuted (Lindsey 2005 ; Nelson et al. 2013 ).
Besides providing space for wild ungulates, small wildlife reserves (with sizes usu-
ally between 200 and 600 km
2 ) at the interface between protected and non-protected
areas have a high potential to provide additional space for lions: when landowners
create collaboratively managed conservancies, land use tend to shift to high-value
trophy hunting and ecotourism where lions and other predators are considered valuable
(Lindsey et al. 2009 ; Nelson et al. 2013 ).
The Etosha Rand Lion Conservation Project
Wildlife–human confl ict along borders between protected and non-protected areas
is one of the main threats facing the African lion. The “Etosha Rand Lion
Conservation Project” in Namibia focuses on providing a scientifi c basis for
improved management of lions at the interface between protected and non-protected
areas. Here, levels of human–lion confl ict are particularly high and the resulting
indiscriminate killing of lions represents one of today’s main threats to the species.
The “Etosha Rand Lion Conservation Project” aims to provide systematic data of
lion population structure and distribution on a wildlife reserve in Namibia located at
the protected/non-protected area interface. Over the last 20 years, more than 1000
lions have been killed by livestock farmers and wildlife reserve managers on lands
bordering Etosha. In other small wildlife reserves in Namibia, culling—similar to
small reserves in South Africa—is often carried out in response to overpopulation
resulting from using electric fences to contain lions within reserve boundaries.
M. Trinkel and F.M. Angelici
59
The “Etosha Rand Lion Conservation Project” is conducted on a wildlife reserve in
Namibia, bordering on Etosha National Park. In contrast to other fully electrifi ed
small wildlife farms, the reserve is separated from the National Park by a semiper-
meable fence through which lions (but not prey species) can freely move and thus
function as part of a much larger ecosystem. However, where the reserve borders
onto livestock farms, an electrifi ed fence has been erected. The “Etosha Rand Lion
Conservation Project” aims to establish essential baseline data on population
dynamics of lions on a reserve with semipermeable fencing and contribute to the
development of a novel management approach for lions in non-protected reserves,
and potentially reduce the extent of indiscriminate killing. The data will help
provide the basis for developing new management approaches of lions outside
protected areas, designed to increase the land area available to the species.
Trophy Hunting of Lions as a Conservation Tool
Trophy hunting can be a useful conservation tool with the potential to increase the
range of African lions , but it can also be a threat, depending on how trophy hunting
is regulated and managed (Whitman et al. 2004 ; Loveridge et al. 2007 ; Packer et al.
2011 ). Hunting of lions for sport is permitted in 23 African countries, with southern
and eastern Africa having the largest hunting industries (Lindsey et al. 2006 ). Trophy
hunting of lions plays a minor role in West and Central Africa (Lindsey et al. 2006 ).
At present, African lions are permitted to be trophy hunted in an area of about
558,000 km
2 (Lindsey et al. 2013 ; Riggio et al. 2013 ). This represents 27–32 % of
the range of the lion in countries where trophy hunting is generally allowed, and at
least 16 % of the total lion distribution range in Africa (Lindsey et al. 2013 ; Riggio
et al. 2013 ).
There are a number of characteristics making the hunting industry suitable to play
an important role in conservation (Baker 1997 ; Bond 2004; Lindsey et al. 2006 ):
Hunters have a lower impact on the environment than ecotourists in terms of
disturbances, conversion of habitat, and the use of fossil fuel.
The infrastructure required for ecotourism is more expensive than that required
for hunting.
The income per hunter is higher compared to the income per tourist. Thus, many
more tourists than hunters are needed to generate the same revenue.
Hunting operators can generate income in areas which are not suitable for eco-
tourism, such as remote areas, areas with low wildlife population densities, areas
where—in addition to wildlife—livestock and local people are present, areas
and/or countries which are politically unstable.
Trophy hunting creates an economic incentive to generate and income from
wildlife in areas that might otherwise be used for agriculture or livestock.
Problems with trophy hunting include unethical hunting practices and overhar-
vesting (Loveridge et al.
2007 ; Packer et al. 2011 ). Unethical hunting practices
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
60
include shooting from vehicles, shooting young individuals and “ canned hunting ,”
where captive-bred lions are hunted in small enclosures, mainly in South Africa
(Damm 2005 ; Lindsey et al. 2006 ). Although executed only by a minor number of
operators (Lindsey et al. 2006 ), such unethical hunting practices have contributed to
a negative publicity of trophy hunting as a conservation tool (Vartan 2002 ). NGOs,
mainly from Europe and North America, have undertaken several attempts to ban
trophy hunting (Lindsey et al. 2013 ). Trade bans can be fatal for the retrieval and
retention of land for wildlife and the tolerance towards lions (Lindsey et al. 2006 ).
Several countries have taken the decision to ban hunting completely. For exam-
ple, in Kenya trophy hunting has been banned since 1977. In that time, wildlife
numbers have declined by 60–70 % since the 1970s, mainly because of a lack of
incentives to invest in wildlife as a form of land use (Norton-Griffi ths 2007 ; Western
et al. 2009 ; Croes et al. 2011 ; Lindsey et al. 2013 ; Nelson et al. 2013 ). Although
ecotourism in Kenya generates large amounts of income, wildlife tourism is con-
ducted in only 5 % of the land, which limits the distribution of income generated by
wildlife (Norton-Griffi ths 2007 ). Similar negative consequences on wildlife num-
bers following hunting bans were observed in Tanzania and Zambia , where sport
hunting was forbidden from 1973 to 1978 and 2000 to 2003, respectively (Baker
1997 ; Leader-Williams and Hutton 2005 ; Lindsey 2005 ; Lewis and Jackson 1997 ).
According to Lindsey et al. ( 2006 ), therefore, avoiding future hunting bans is vital
for lion conservation.
Lions are one of the most economically valuable species in Africa’s trophy hunting
industry and are therefore most prone to over-harvesting (Lindsey et al. 2012b ).
In fact, recent studies demonstrated that over-harvesting may have contributed to lion
declines in some areas in Tanzania (Packer et al. 2009 , 2011 ). Furthermore, hunting
activities in non-protected areas were found to extend into the unhunted lion popula-
tion inside the protected area (Loveridge et al. 2007 ): excessive trophy hunting
changed the demography of the lion population in Hwange National Park, Zimbabwe,
although not reducing the viability of this lion population (Loveridge et al. 2007 ).
Other areas of Zimbabwe have not faired so well, with lions in Gonarezhou National
Park and Tuli Safari Area both suffering a collapse of numbers due to excessive trophy
hunting (Groom et al. 2014 ). Groom et al. ( 2014 ) do recognize the importance of
trophy hunting in the conservation of the lion; however, they call for stricter regula-
tions to ensure the long-term survival of lions in hunted areas.
Whitman et al. ( 2004 ) suggested that implementing a 6-year age minimum would
ensure safe harvest irrespective of population size. Some countries have made
efforts to make lion hunting more sustainable in recent years and off-takes have
declined signifi cantly (Lindsey et al. 2013 ). For example, Tanzania and
Mozambique’s Niassa Reserve have successfully implemented a 6-year age mini-
mum for hunted lions (Begg and Begg 2009 ; Packer et al. 2011 ). However, in many
countries improved management of lion hunting to prevent negative impacts on
hunted lion populations is still lacking (Lindsey et al. 2013 ). According to Loveridge
et al. ( 2007 ) and Lindsey et al. ( 2013 ), main changes needed are as follows: (1) a
reduction in quotas to realistic levels (no more than 10 % of adult males) based on
robust population estimates in some countries, (2) an implementation of trophy
monitoring and adequate quota management, (3) enforcing an age minimum with
M. Trinkel and F.M. Angelici
61
appropriate penalties for harvesting animals below a threshold age in all countries,
and (4) a lion hunt should have a minimum length of 21 days. As lions in West and
Central Africa are Critically Endangered, hunting, although playing a minor role in
these areas, should be strictly prohibited.
Concluding Remarks
African lions have lost most of their historical range and continue to decline. Lion
populations in West and Central Africa and in some parts of East Africa show an
alarming downward trend which presses the need for urgent conservation actions.
In West and Central Africa , and in Ethiopia, where remaining lion populations
are extremely small and isolated, conservation measures have to be taken quickly.
Besides mitigating human–lion confl icts at reserve borders, the security of protected
areas needs to be increased (e.g., by effi cient prevention of poaching of lions and
their prey) and protected areas should be connected through ecological corridors to
maintain genetically viable populations. Failing this (or while it is being imple-
mented), translocation schemes may be needed in the short term to conserve or
increase genetic diversity. Hunting, although playing a minor role in West and
Central Africa, should be strictly prohibited in protected areas and areas forming
wildlife corridors. To counteract inbreeding of the Critically Endangered population
in West and Central Africa, it might be essential to initiate a captive breeding pro-
gram (ex situ) (Barnett et al. 2006) to increase the size of the population and its
genetic diversity. In areas, where the species has disappeared and which meet the
necessary ecological conditions (e.g., prey availability, protection from poaching),
reintroduction of prey should be considered.
Confl ict mitigation between humans and wildlife can be very effective and small
changes in livestock husbandry can successfully reduce livestock losses and subse-
quently reduced the number of lions killed in many areas in East and West Africa.
When developing such techniques, the tradition and culture of local communities
have to be taken into consideration, and methods that proved to be successful in one
area may not necessarily be applicable to other sites. Other approaches for effective
long-term conservation of African lions favor the separation of land used by humans
and conservation areas through the erection of fences. Fences have already been
erected in many areas in southern Africa, where lion populations are still viable. In
southern Africa and some parts of East Africa, where lions are still abundant, eco-
tourism and hunting have been successfully implemented to stabilize lion popula-
tions. To prevent negative impacts on hunted lion populations, however, improved
management of lion hunting is necessary in many countries.
Since African lions occur in different macro-areas, i.e., in West and Central
Africa, in the Ethiopian district, in East Africa and in southern Africa, these areas
should be precisely re-defi ned geographically and conservation strategies for lions
should be adjusted according to these geographical differences, based on a probable
redefi nition of the taxonomy of Panthera leo . This may result in different P. leo
subspecies with the need to adjust conservation actions within an overall strategy
3 The Decline in the Lion Population in Africa and Possible Mitigation Measures
62
for the conservation of the species as a whole (Walston et al. 2010 ). Such an overall
conservation action plan should also include the allocation of funds by involved
countries, international institutions, and private donors. For example, for the tiger
Panthera tigris with its different populations and subspecies, a multiannual program
was drawn up. The so-called Global Tiger Recovery Program was approved by all
countries in which the tiger occurs and aims to achieve the objectives (e.g., fi ght
against poaching or trade in body parts, regular monitoring of tiger populations,
increasing the protection of tigers in both protected and non-protected areas) within
12 years (2010–2022) (World Bank 2011 ).
There are many factors threatening lions, which vary between sites. Multiple
inventions will often be required and the suitability of adequate solutions will differ
from area to area. Failure to address the problem will have fatal consequences for
the survival of lions and wildlife in general.
Acknowledgments We wish to thank Matt W. Hayward, Susan M. Miller, and Craig Packer for
comments and suggestions on an earlier draft which considerably improved this chapter. Lorenzo
Rossi provided the fi gure.
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M. Trinkel and F.M. Angelici
... Livestock frequently appear in the diets of in big cats across the globe (Ghouddousi et al., 2016;Babrgir et al., 2017). Local farmers may opt to poison or shoot big cats in order to protect their livestock (Trinkel et al., 2016). In some areas, direct persecution may represent a large proportion of felid mortalities. ...
... In some areas, direct persecution may represent a large proportion of felid mortalities. For example, Trinkel et al. (2016) found that within their study period human-wildlife conflict caused almost 50% of lion mortalities on the border of Etosha National Park. Subadult males and adult females represented the majority of persecuted individuals, and this had second-order effects on the population structure of lion prides (Trinkel et al., 2016). ...
... For example, Trinkel et al. (2016) found that within their study period human-wildlife conflict caused almost 50% of lion mortalities on the border of Etosha National Park. Subadult males and adult females represented the majority of persecuted individuals, and this had second-order effects on the population structure of lion prides (Trinkel et al., 2016). Moreover, Tortato et al. (2017) suggest that depredation of large livestock may lead to a greater risk of infanticide due to attraction of multiple individuals to a carcass, including females with cubs. ...
Thesis
We live in a world of human-induced rapid environmental change, where the frequency of extinctions and resulting loss in biodiversity has reached levels associated with a mass extinction event. At the same time, technological developments in computing have facilitated the growth of highly complex, mechanistic models across all scientific fields. The challenge for conservation biologists is then to develop models that can predict how organisms respond to conservation measures and increasing anthropogenic pressures. Here I explore the potential and limitations for conservation applications of spatially-explicit mechanistic models of habitat selection, by developing a simulation applicable to large felids. I demonstrate that initial choice of resolution may bias the parameterisation process of spatially-explicit models, when applied to spatially-explicit empirical data. I use mechanistic models to address two current problems in conservation biology: (a) efficient calculation of movement metrics from telemetry data, tested with a virtual ecology approach; and (b) accounting for interacting influences on populations, quantified with a model that controls for confounding variables. I identify the major caveats to accurately predicting the complex behaviour of large-bodied animals. The spatially-explicit mechanistic models developed here, and applied to real-world problems, demonstrate the potential of these types of simulation for confronting otherwise impossible questions in diverse areas of conservation biology.
... Despite decades of applied conflict management research (Trinkel and Angelici, 2016;van Eeden et al., 2018), sustainable coexistence of rural communities with lions has yet to be achieved in many countries (Bauer et al., 2015). The successful mitigation of conflict primarily depends on changes in people's behaviors and risk management (Reddy et al., 2017), requiring trans-disciplinary research and conservation approaches that appropriately reflect the human dimensions of coexistence (Bennett et al., 2017;Pooley et al., 2017). ...
... In northern Botswana, communities seek safety from lions, primarily via physical separation and lion control. However, the continued lethal removal of damagecausing lions threatens population viability Trinkel et al., 2017); fencing is expensive (Packer et al., 2013), ecologically problematic (Trinkel and Angelici, 2016), and can shift problems elsewhere (Osipova et al., 2018); while translocations are resource-intensive and have historically shown low success rates (Stander, 1990). By default, separation undermines coexistence. ...
Article
Full-text available
Across Africa, lions (Panthera leo) are heavily persecuted in anthropogenic landscapes. Trans-disciplinary research and virtual boundaries (geofences) programmed into GPS-tracking transmitters offer new opportunities to improve coexistence. During a 24-month pilot study (2016-2018), we alerted communities about approaching lions, issuing 1,017 alerts to four villages and 19 cattle posts. Alerts reflected geofence breaches of nine lions (2,941 monitoring days) moving between Botswana's Okavango Delta and adjacent agro-pastoral communities. Daily alert system costs per lion were US$18.54, or $5,460.24 per GPS deployment (n = 13). Alert-responsive livestock owners mainly responded by night-kraaling of cattle (68.9%), significantly reducing their losses (by $124.61 annually), whereas losses of control group and non-responsive livestock owners remained high ($317.93 annually). Community satisfaction with alerts (91.8%) was higher than for compensation of losses (24.3%). Study lions spent 26.3% of time monitored in geofenced community areas, but accounted for 31.0% of conflict. Manual alert distribution proved challenging, static geofences did not appropriately reflect human safety or the environment's strong seasonality that influenced cattle predation risk, and tracking units with on-board alert functions often failed or under-recorded geofence breaches by 27.9%. These insufficiencies prompted the design of a versatile and autonomous lion alert platform with automated, dynamic geofencing. We co-designed this prototype platform with community input, thereby incorporating user feedback. We outline a flexible approach that recognizes conflict complexity and user community heterogeneity. Here, we describe the evolution of an innovative Information and Communication Technologies-based (ICT) alert system that enables instant data processing and community participation through interactive interfaces on different devices. We highlight the importance of a trans-disciplinary co-design and Weise et al. Lion Early Warning Botswana development process focussing on community engagement while synthesizing expertise from ethnography, ecology, and socio-informatics. We discuss the bio-geographic, social, and technological variables that influence alert system efficacy and outline opportunities for wider application in promoting coexistence and conservation.
... Acceptance of a two-subspecies arrangement largely based on some genetic data seems to overly neglect the great phenotypic diversity still found, for instance, in lion Panthera leo (Linnaeus, 1758) [143,144] and ignores some studies that highlight genetic divergence promoted by ecological discontinuity [145,146] that seems to support a new paradigm to explain microevolutionary divergence in widespread, large-sized carnivores [147]. Therefore, without forgetting or underestimating the composite problems of lion conservation, and of coexistence/interaction with humans involving this highly charismatic species [148,149], taxonomic issues deserve to be included in the future conservation strategies of this charismatic species. Paradoxically, tourists (but also trophy hunters) may be greatly interested in phenotypic lion diversity, and works such as [143] may provide the input for further travels in different regions of Africa and the promotion of new conservation/tourism projects for littleknown, overlooked populations such as those of Southwestern Ethiopia. ...
Article
Full-text available
Ecotourism can fuel an important source of financial income for African countries and can therefore help biodiversity policies in the continent. Translocations can be a powerful tool to spread economic benefits among countries and communities; yet, to be positive for biodiversity conservation, they require a basic knowledge of conservation units through appropriate taxonomic research. This is not always the case, as taxonomy was considered an outdated discipline for almost a century, and some plurality in taxonomic approaches is incorrectly considered as a disadvantage for conservation work. As an example, diversity of the genus Giraffa and its recent taxonomic history illustrate the importance of such knowledge for a sound conservation policy that includes translocations. We argue that a fine-grained conservation perspective that prioritizes all remaining populations along the Nile Basin is needed. Translocations are important tools for giraffe diversity conservation, but more discussion is needed, especially for moving new giraffes to regions where the autochthonous taxa/populations are no longer existent. As the current discussion about the giraffe taxonomy is too focused on the number of giraffe species, we argue that the plurality of taxonomic and conservation approaches might be beneficial, i.e., for defining the number of units requiring separate management using a (majority) consensus across different concepts (e.g., MU-management unit, ESU-evolutionary significant unit, and ECU-elemental conservation unit). The taxonomically sensitive translocation policy/strategy would be important for the preservation of current diversity, while also supporting the ecological restoration of some regions within rewilding. A summary table of the main translocation operations of African mammals that have underlying problems is included. Therefore, we call for increased attention toward the tax-onomy of African mammals not only as the basis for sound conservation but also as a further opportunity to enlarge the geographic scope of ecotourism in Africa.
... Direct interactions among predators, humans and human assets such as livestock are the most pronounced causes of human-wildlife conflicts (e.g. Haque et al. 2015;Olson et al. 2015;Trinkel and Angelici 2016). Human-wildlife conflict can also come about through impacts of predators on high-value conservation assets such as populations of threatened species (e.g. ...
Article
Context Exclusion fences are increasingly used to prevent interactions between predators (introduced and native) and assets such as endangered species or livestock. However, challenges remain in identifying when exclusion fences are an optimal investment and the intended outcome is likely to be achieved. The level of association with complementary methods of control that is needed is also unclear. AimsWe aimed to quantify the interactions among factors that affect fencing efficiency, including the size of the fenced area, the fence permeability, the initial density of the predator population, and its survival of complementary control methods. Methods Using a spatially explicit, individual-based model, we simulated wild dog (dingo) populations as a proxy for describing predator dynamics inside a fenced area under different management practices and fence designs. We then fit a generalised linear model to the model outcomes to assess the effects of the four factors mentioned above. Key resultsLethal control had a strong effect on wild dog density when the survival of control was lower than 0.5. Fences generally had an effect on wild dog density only when their permeability was lower than ~1% and their effect was most noticeable when the initial density was very low (<2 dogs per 100km2), or when survival of control was very low (<0.5). Conversely, when the initial density was very high (~12 dogs per 100km2), a fence with a low permeability (<1.5%) caused the paradoxical effect that wild dog density could be higher than that obtained with a more permeable fence. Wild dog eradication was possible only when survival of control was 0.25 or lower, except when either initial density or fence permeability were extremely low (<2 dogs per 100km2 and <0.1% respectively). Conclusions Our results demonstrated that large exclusion fences can be an effective aid in managing predator populations. We recommend that this tool should be used as a preventive measure before predators establish a population inside the area targeted for exclusion, in tandem with lethal control, or when an initial marked reduction of predator density can be achieved. We also demonstrated that eradication can be achieved only when a narrow combination of parameters is met. Implications Land managers should carefully evaluate when and at what scale control tools should be deployed to control wild dog populations. Landscape application of exclusion fences faces the challenge of high maintenance to ensure low permeability, coupled with very high sustained suppression of wild dog density, which are unlikely to be feasible options in the long term. Conversely, the same control techniques could provide efficient asset protection at a smaller scale where fence maintenance and sufficient control effort can be sustained.
... Human-wildlife conflicts are one of the frequent challenges faced by managers around the world (Madden 2004). Of these conflicts, direct interactions between predators and humans, or human assets such as livestock, are of greatest public concern (Haque et al. 2015;Olson et al. 2015;Trinkel and Angelici 2016). A range of tools is employed to mitigate these impacts and population models are often used to improve planning efforts and test possible intervention strategies (Himes Boor 2014). ...
Article
ContextManagement of human–wildlife conflicts is of critical importance for both wildlife conservation and agricultural production. Population models are commonly used to simulate population dynamics and their responses to management actions. However, it is essential that this class of models captures the drivers and mechanisms necessary to reliably forecast future system dynamics. AimsWe aimed to develop a flexible modelling framework with the capacity to explicitly simulate individual interactions with baits (with or without the presence of other management tools), for which parameter estimates from field data are available. We also intended for the model to potentially accommodate multi-species interaction and avoidance behaviours. Methods We expanded an existing spatially explicit, individual-based model to directly simulate bait deployment, animal movements and bait consumption. We demonstrated the utility of this model using a case study from Western Australia where we considered two possible exclusion-fence scenarios, namely, the completion of a landscape-scale and smaller-scale fences. Within each of these proposed cells, using data obtained from a camera-trap study, we evaluated the performance of two levels of baiting to control wild dogs (Canis familiaris), in contrast with the option of no control. ResultsThe present study represents a substantial step forward in accurately modelling predator dynamics. When applying our model to the case study, for example, it was straightforward to investigate whether outcomes were sensitive to the bait-encounter probability. We could further explore interactions between baiting regimes and different fence designs and demonstrate how wild dog eradication could be achieved in the smaller cell under the more intense control scenarios. In contrast, the landscape-scale fence had only minor effects unless it was implemented as a preventive measure in an area where wild dogs were not already established. Conclusions The new component of the model presented here provides fine-scale control of single components of individual–bait interactions. ImplicationsThe effect of management actions (e.g. lures) that affect this process can be easily investigated. Multi-species modelling and avoidance behaviours can readily be implemented, making the present study widely relevant for a range of contexts such as multi-species competition or non-target bait uptake.
... Rajaratnam et al., 2016;Penteriani et al., 2017;Moreira-Arce et al., 2018;Kelly et al., 2019;Ugarte et al., 2019;Reyna-Saenz et al., 2020) and the ensuing persecution and threats to several species and/or local populations (e.g. Bauer et al., 2015;Jacobson et al., 2016;Trinkel & Angelici, 2016;Durant et al., 2017). Recovery of large carnivore populations and co-existence with humans, however, are not impossible, as exemplified by the current situation in Europe with grey wolves and brown bears (Chapron et al., 2014). ...
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Small carnivores – here defined as members of the mammalian Order Carnivora with a body mass < 21.5 kg – occur worldwide, including in Oceania, following introductions. They are represented by 210 to 282 species, which corresponds to around 90% of terrestrial carnivores globally. Some species are endemic to one or two countries (sometimes only islands), while others, like the red fox, Vulpes vulpes, are present in nearly 90 countries over five continents. Small carnivores inhabit virtually all of the Earth’s ecosystems, adopting terrestrial, semi-fossorial, (semi-)arboreal or (semi-)aquatic lifestyles. They occupy multiple trophic levels, being primary consumers when feeding on fruits, seeds, and other plant matter, secondary consumers when preying on frugivorous, granivorous, and herbivorous animals, or tertiary consumers when killing and devouring meat-eating animals. Therefore, they play important roles in the regulation of ecosystems, e.g. natural pest control, seed dispersal and nutrient cycling. In areas where humans have extirpated large carnivores, small carnivores may become the dominant predators, which may increase their abundance (‘mesopredator release’) to the point that they can sometimes destabilize communities, drive local extirpations, and reduce overall biodiversity. On the other hand, one-third of the world’s small carnivores are Threatened or Near Threatened with extinction (sensu IUCN). This results from regionally burgeoning human populations’ industrial and agricultural activities, causing habitat reduction, destruction, fragmentation, and pollution. Overexploitation, persecution, and the impacts of introduced predators, competitors, and pathogens have also negatively affected many small carnivore species. Although small carnivores have been intensively studied over the past decades, bibliometric studies showed that they have not received the same attention given to large carnivores. Furthermore, there is a huge disparity in how research efforts on small carnivores have been distributed, with some species intensively studied, and others superficially or not at all. Regionally, North American and European small carnivores have been the focus of numerous studies, and more research is being progressively conducted in Asia. However, there is a need to increase the research effort in Africa and Central and South America. Encouragingly, the recognition of the importance of the mesopredator release effect and the exponential deployment of camera-traps have started to boost the research effort and scientific knowledge on small carnivores around the world. This book aims at filling a gap in the scientific literature by elucidating the important roles of, and documenting the latest knowledge on, the world’s small carnivores. It is divided into four main sections: (i) Evolution, Systematics, and Distribution; (ii) Ecology, Behaviour, and Diseases; (iii) Interspecific Interactions and Community Ecology; and (iv) Interactions with People and Conservation. We hope that the book will appeal to a wide audience and, considering that the field of small carnivore science remains wide open, stimulate much-needed research globally.
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An important pillar to the mainstream conservation of elephants (Loxodonta africana, L. cyclotis and Elephas maximus) is the relatively narrow and growing interest of ‘human-elephant conflict’ prevention and mitigation. This thesis problematises the hegemonic HEC discourse (Igoe, Neves and Brockington, 2010:488) which acts as a powerful resource to promote human-elephant separation, often producing failing and harmful practices, which are often obscured from public view. The discourse frames human-elephant interactions as conflict with elephants, blames local population growth and encroachment for its escalation and in turn fixates on technocratic methods of separation and control while promoting many projects as ‘win-win’ . As methods of separation are favoured, compensation is routinely tested but derided and local financial resilience, autonomy and values largely ignored. Methods of separation are replicated across Asia and Africa, finding some success to avoid crop loss. However, separation seems ill-suited to elephants, ecologically dangerous, physically difficult, and unevenly costly. It is also an appeasement to agribusiness , an acceleration of neoliberal capitalism within traditional communities, a reprise of fortress conservation and a tool to legitimise harmful forms of nature’s commodification. These issues highlight the need to question human-elephant separation and awaken opportunities to realise actively shared spaces.
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In lion conservation, the International Union for Conservation of Nature (IUCN), the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), and Trade Records Analyses of Flora and Fauna in Commerce (TRAFFIC) are considered key United Nations (UN) institutions for “science-based decisions” on global policy formulation for conservation and combating wildlife crime. CITES, but probably also TRAFFIC and IUCN, still adheres to and operates in the paradigm of the “sustainable use” of animals, based on the long leading philosophical Cartesian paradigm in academia that premises that humans and animals differ in kind, and that animals do not “feel” and have the neurological capacities to think like us. But this Cartesian worldview can no longer withstand the latest scientific evidence, developments, and new insights that show how people and animals only differ in degree and not in kind. The concept of “sustainable use” of wildlife, including lions, therefore needs to be rethought in the light of this new paradigm. In South Africa, the “sustainable use” of lions includes the trade in lion’s bones from captive lions, which was legalized in 2016. The Wildlife Animal Protection Forum of South Africa (WAPFSA) appealed against this legalized trade, based on rational arguments that fit CITES and its Cartesian approach to animals but also on the paradigm shift where humans and animals are no longer considered different in kind but only in degree. This paradigm shift has led to initiatives to try and suggest possible ways forward for a political order that matches this “new normal.” Probably the most developed in this context is the concept of “zoopolis,” which is explored in this article. The four “vulnerabilities” on which the concept is based were all found to be relevant to lion conservation and fighting wildlife crime in South Africa.
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The vanishing population of big cats is a global problem difficult to solve. In modern era several subspecies and populations of felids of the genus Panthera have become extinct, or locally extinct. The review of their story seems to show a common trend: a decrease of prey due to hunting and habitat destruction that leads to the decrease of predators, which switch to livestock creating conflicts with human. In many cases the micro populations of big cats are very difficult to detect, surviving for several years after their official extinction without it being possible to implement conservation strategies. To be effective, conservation measures must be as interdisciplinary as possible and include the active involvement of the locals.
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Wherever wildlife management concerns the movement of individuals across structured habitat, its scale of operations will encompass metapopulation dynamics. The goal of this chapter is to review the potential applications of metapopulation concepts and models in reserve design and conservation management. Our perspective is forward-looking. We show how some key problems of where to direct conservation effort and how to manage populations can be addressed in the context of regional habitat structure and the survival and renewal of habitat patches. We also mention several cases of successful metapopulation management and point out practical problems. We emphasise (1) that the viability of a population may depend on surrounding populations, in which case metapopulation processes influence or determine reserve design and management options; (2) that understanding the dynamic processes requires models, which make assumptions that need validating; (3) that the principle limitation of metapopulation models is their single-species focus. Conservation strategies clearly depend on the particular social, economic and ecological circumstances of each region, and concepts such as the metapopulation can seem irrelevant to practical concerns. We aim to show, nevertheless, that an understanding of metapopulation dynamics can be vital to asking pertinent questions and seeking potential solutions. The conceptual framework of metapopulation dynamics tells us what information is needed in order to build case-specific models relevant to any of a wide range of issues. These issues include the potential disadvantages of habitat corridors, or hidden benefits of sink habitat; the optimal schedule for translocations or reintroductions; the relative merits of reducing local extinctions against increasing colonisations; the optimum distribution of habitat improvement; and the advantages of increasing life spans of ephemeral habitats.
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The late Pleistocene vertebrate record of Africa contributes to our understanding of human evolution and the development of modern biotic environments. This article explores this record, paying particular attention to terrestrial mammals. Because the origins of physically and behaviorally modern humans occurred in Africa during the late Pleistocene, much of what we know about vertebrates from this time comes from archaeological research. Most of the mammal species found in Africa during the late Pleistocene were also found here historically; only a few African species went extinct during the late Pleistocene. However, many changes in species’ ranges, abundance, and associations made late Pleistocene faunal communities look different from today's communities. As climate changed, so did vegetation, creating unique habitats, many of which do not exist today. Each species’ unique responses to these changes caused the formation of mammal communities that do not have analogs today, meaning that species are often found together in fossil assemblages that were not found together historically. Often fossil species are found in places where today their descendants live hundreds of kilometers away, indicating that their past ranges had either shifted or expanded.