Review: Tool or Threat: Is trophy hunting of Lions of Conservation Value?

ARTICLE INFO Keywords:

Panthera Leo Trophy hunting Sub-Saharan Africa Sustainability Trade offs Conservation values

ABSTRACT Trophy hunting has long been associated with conservation, with many western conservation policies stemming from colonial hunting especially in sub-Saharan Africa. As a highly emotive topic, it has drawn great debate in recent times, with polarized arguments either heralding it as an important and necessary conservation tool, or attacking it as a grotesque, unsustainable and deleterious practice. This paper discusses the conservation role of trophy hunting through an evaluation of ecological evidence and human values. This paper focuses predominantly on its significance as both a tool and threat to African lion conservation, but its principles will apply for other felids. Trophy hunting of lions stimulates protection of habitat and maintenance of sustainable populations in areas where there is no conservation alternative. However, there is little evidence of human and ecological conservation benefits, and the practice is replete with poor management and corruption. The potential of trophy hunting as a conservation tool is context/taxa specific. Its value may be considered in terms of the conservation of species, populations or individuals. Each level can in turn be considered in terms of its contrasting instrumental and intrinsic value. Trophy hunting is thus both a tool and threat to lion conservation. Ultimately, this discussion accepts that regardless of the scales of analysis used, or the apparent practical and theoretical issues surrounding it, trophy hunting is here to stay. The practical, not theoretical, debate is thus not whether trophy hunting is a conservation tool, but whether it can be improved so that animals and humans can benefit as much as possible from this polarizing practice.

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Contents

Abbreviations  .....................................................................................................................  2  

1   Introduction:  paradoxical  conservation  .............................................................  3  2   Trophy  Hunting  of  lions:  Conservation  tool  ......................................................  6  2.1   Benefits  of  trophy  hunting  ...........................................................................................  6  2.2   Alternative  conservation  ..............................................................................................  7  2.3   Sustainability  ....................................................................................................................  8  

3   Trophy  hunting  of  lions:  Conservation  threat  ...............................................  10  3.1   Limitations  and  vulnerability  ...................................................................................  10  3.2   Deleterious  impacts  ......................................................................................................  12  

4   Conservation  Values  ...............................................................................................  15  4.1   Intrinsic-­‐Instrumental  Trade  offs  ............................................................................  16  

5   Conclusion  ..................................................................................................................  18  6   References  ..................................................................................................................  19  

7   Figures  .........................................................................................................................  25  

Abbreviations  IUCN International Union for Conservation of Nature PA Protected Area PETA People for the Ethical Treatment of Animals

List  of  figures  Fig. 1: Number Foreign hunters in sub-Saharan Africa………………………..…..P 26 Fig. 2: Proportion of hunters visiting each country in sub-Saharan Africa.……….P 26 Fig. 3: Effects of 30 years of lion trophy hunting as a function of hunting quota size and male age in a hypothetical population………………………………………... P 27 Fig. 4: Age estimation for adult lions using nose colouration..………………...…P 27 Fig. 5: Effects of 40 years of trophy hunting of male lions on female population size as a function of hunting quota size and male age ……………………………….…P28 Fig. 6: Stochastic model using 40 years of trophy hunting data from the Serengeti…………………………………………………………………………….P28 Fig. 7: Boundaries of different types of protected areas in Tanzania………………P29 Fig. 8: Average number of lions and leopards harvested in major hunting areas..…P29 Fig. 9: Estimated proportion of the lion population in each country removed annually by trophy hunting ………………………………………………..….………….....P30

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1 Introduction:  paradoxical  conservation     Trophy hunting is an important, emotive and controversial aspect of conservation

(Palazy et al., 2011). The controversy derives from the fact that it can be both a

significant threat to and a tool in conservation, especially of African lions. Indeed, its

aims, including sustainable use of natural resources, high economic benefits and low

environmental impacts (Loveridge et al., 2006), make trophy hunting a potentially

important conservation tool (Hutton and Leader-Williams, 2003; Lindsey et al.,

2007). However this view is challenged by the argument that the negative effects of

poor governance and unsustainable practices outweigh its ecological and economic

benefits. Trophy hunting is considered by some to be a significant threat to

endangered species (e.g. Courchamp et al., 2006), and is also vehemently opposed

from an ethical standpoint (People for the Ethical Treatment of Animals [PETA],

2012).

Contemporary Western conservation, especially the policy of protected areas (PA),

has its roots in nineteenth century colonial trophy hunting (Adams, 2004), where wild

species were protected in order to be hunted by wealthy visitors (Loveridge et al.,

2006). Despite significant populist condemnation (e.g. Laing, 2012), trophy hunting

remains a popular practice (Figs 1 & 2) across North America and in sub-Saharan

Africa, where over 1.3million km2 of land is used for hunting. This exceeds the area

encompassed by national parks by 22% in the countries where hunting is permitted

(Lindsey et al., 2007). Trophy hunting generates gross revenues of over US$201

million annually in sub-Saharan Africa, although over half accrues in South Africa

and towards its canned lion hunting industry (Lindsey et al., 2007). Trophy hunting is

important for both conservation and development, as exemplified by its prominent

role in the proliferation of Community-Based Natural Resource Management. Trophy

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hunting is an aspect of ‘sustainable use’ in the Convention on Biological Diversity’s

report (2003) (Loveridge et al., 2006; Bélair et al., 2010), and is increasingly accepted

in the contemporary conservation paradigm of human use/consumption of wildlife.

The growing integration of humans in conservation strategies has generated a need for

trade-offs between human and ecological values. Trophy hunting has therefore to be

framed in terms of conservation values, morals and ethics.

African lions (Panthera Leo) are currently categorized by the International Union for

Conservation of Nature (IUCN) as “vulnerable” (Bauer et al, 2008). Numbers appear

to have declined from 100,000 in 1990 (Nowell and Jackson, 1996) to as low as

23,000 today (Myers, 1975; Chardonnet, 2002; Bauer and van der Merwe, 2004), and

there is concern for the survival of the species. The main threats to lions are illegal

killings (poaching or retaliatory), habitat loss/fragmentation and disease (Milner-

Gulland et al. 2003; Whitman et al. 2004; Ray et al., 2005; Bauer et al., 2008; Caro et

al. 2009; Fryxell et al. 2010). Trophy hunting can also be a threat, with significant

impacts on the viability of populations, especially when seen in the light of the

increasingly alarming context (lower genetic diversity, smaller populations, human-

lion conflict) within which lions now exist. Lions are very important to the trophy

hunting industry (e.g. Lindsey et al., 2012), and they are often the most valuable

species sold in safari hunting concessions (Loveridge et al. 2007). The large home

range of lions and other ecological features pre-dispose them for conflict with human

(Lichtenfeld, 2005). Trophy hunting potentially relieves this conflict and is arguably

paradoxically in the species’ best interest. The recent (partly UK funded) LionAid

conference, where wildlife delegates across sub-Saharan Africa collaboratively

considered future conservation programs and the role of hunting for the survival of

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African lions, is an important step in addressing key issues of management and wider

scale approaches (LionAid, 2012).

This paper reviews data on the influence of trophy hunting on populations of lions. It

examines the ecological and human criteria necessary for sustainable trophy hunting,

and reviews how trophy hunting needs to be evaluated on the basis of viable

alternatives and the necessity for tradeoffs. These are contextualized in the wider

conservation discourse of intrinsic-instrumental values. It proposes that through

careful management, a practical and sustainable line must be drawn between the

opposing paradoxical forces of threat/conservation.

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2 Trophy  Hunting  of  lions:  Conservation  tool  

2.1 Benefits  of  trophy  hunting  

Trophy hunting of lions can be a conservation tool through the process of

incentivisation. Perhaps the most important manifestation of this incentivisation is the

protection and/or acquisition of land (Jew and Bonnington, 2011; Lindsey et al.,

2007; Loveridge et al., 2006). In Zimbabwe, which has seen extensive land

conversions, there are examples of species recoveries/re-introductions though land

protection (Loveridge et al., 2006, 2007; Bond et al., 2004). There are good examples

of species recoveries through trophy hunting (white rhino- Leader-Williams et al.,

2005; leopard- Packer et al. 2009; black wildebeest- Lindsey et al., 2007). Trophy

hunting can also serve to reduce illegal hunting and improve population viability

through the operator’s self-interest in preservation (Lindsey, 2008; Lindsey et al.,

2007, 2012; Leader-Williams and Hutton, 2005). Whitman et al. (2007) note for

example how the removal of wire snares in hunting areas has reduced one

anthropogenic threat to lions. In Zambia, one of the achievements of Administrative

Management Design has been the use of hunting revenues to employ 500 village

scouts for anti-poaching in Game Management Areas (Lewis and Alpert, 1997).

With large home ranges (at least varying between 22 km2 to 7337 km2 in Cameroon

[Van Rijssel et al., 2008]), lions ideally require large tracts of connected habitat, and

this is where trophy hunting could play a significant conservation role on a large

spatial and long-term temporal scale. There is a growing acknowledgement for the

need for conservation action outside of PAs due to habitat loss/fragmentation (Fjeldsa

et al., 2004; Grunblatt et al., 1995; Wilkie and Carpenter, 1999; Willis et al., 2012).

Given trophy hunting’s potential to exist outside of PA’s and incentivize matrix

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biodiversity conservation (especially considering qualitative research suggesting

hunters prefer “unfenced wilderness areas” [Lindsey et al., 2006:288]), the colonial

justification of habitat protection through trophy hunting (see Jepson and Whittaker,

2002), could be one answer. Indeed, perhaps with extensive post-normal (Funtowicz

and Ravetz, 1992) strategic planning with scientists, local communities, hunting

organisations and politicians, combined with government intervention and

regulations, a conscious organization of connected habitats with corridors would

provide a guaranteed (spatial and genetic) sustainable base for a wider network of lion

populations.

Perhaps the most significant trophy hunting benefit is development, although even

this is commonly undermined by elite capture and corruption, whereby the benefits

actually reaching communities are minuscule compared to the total profits

(Loveridge, 2011; Loveridge et al., 2006). Despite the fundamental importance of the

conservation-development intersection, an in-depth exploration of trophy hunting’s

role in socio-economic development will not be pursued here. So, ecologically

speaking, when certain criteria and conditions are met, lion trophy hunting has the

potential to benefit both lions and wider biodiversity. Importantly though, there is a

distinct lack of quantifiable ecological evidence that lions have actually benefitted

from trophy hunting (Lindsey et al., 2007; Loveridge et al., 2006; Packer et al. 2011).

 

2.2 Alternative  conservation  

Moreover, given the ways in which trophy hunting can act as a conservation tool,

perhaps the strongest argument in favour of trophy hunting is that it provides the best

conservation alternative in many instances. Photographic tourism generates up to

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twice the revenue of trophy hunting (Whitman et al., 2007; Loveridge et al., 2006).

However, in areas unsuitable or unused for wildlife purposes like photographic

tourism, converting them into wildlife (hunting) areas is arguably better than its prior

non-consumptive use. Jenkins et al. (2003) allude to Tanzania’s remote and less

scenically attractive Kilombero Game Controlled Area, where trophy hunting has

provided benefits such as protection against poaching and livestock encroachment.

Furthermore, multiple authors note the resilience of the trophy hunting industry to

political instability, especially in the hunting rich country of Zimbabwe (Leader-

Williams and Hutton, 2005); there, hunting revenues dropped by only 12% (compared

to the tourism industry’s 75%) in the first year of the land seizures (Booth, 2002;

Bond et al., 2004). Lindsey et al., (2006, 2007) argue that trophy hunting is a

complementary consumptive resource use, and can therefore be used as part of a

wider conservation strategy, such as allowing hunting only during non-tourist seasons

or at very specific narrow times of the day (Whitman, 2002). Thus, despite significant

limitations that undermine the viability of populations, the potential for wildlife

habitat acquisition/protection and lion population sustainability, means that trophy

hunting can be pragmatically employed as a conservation strategy in areas unsuitable

for photographic tourism.

2.3 Sustainability  

Importantly, trophy hunting of lions has the potential to act in a sustainable manner,

and thereby provide wider aforementioned conservation benefits. The two main ways

of ensuring this are through a minimum age and strict quotas. Since the age of the lion

killed is so ecologically significant, if an age-based criterion to trophy selection is

applied (at least 6 years old), Whitman et al. (2004, 2007) argue that trophy hunting

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can be biologically sustainable- even without the use of quotas (Fig. 3). Following

“[Craig] Packer’s black nose theory” (Damm, 2008:8), Whitman et al. (2004)

strongly support the use of nose pigmentation for estimating age (Figs 4 & 5),

demonstrating how an age-based criterion can be successfully applied. In practice,

since age restrictions have not, and probably will not be, followed 100%, enforcing

quotas is still vital in reducing potential deleterious consequences. Indeed, others

advocate the need for varying degrees of quotas, which must of course be context

dependent (Packer et al., 2011). Although Tanzania’s annual off takes are lower than

its quota of 500 (average of 243 between 1996 and 2006), Lindsey et al. (2012) and

Packer et al. (2011) recommend, in addition to a minimum 6 year age limit, maximum

off takes of 0.5 per 1000 km2 across Tanzania, with an increase to 1.0 in the Selous

game reserve.

The potential to achieve a sustainable balance is further substantiated by evidence

suggesting the resilience of trophy selection to environmental disturbance through

stochastic modeling (Fig. 6) (Whitman et al., 2007). The social stability of lions

appears to increase their resilience to population collapse (Packer et al. 2006). Thus,

given few clear-cut cases of genetic, behavioural and population impacts having

significant impacts on the viability of populations (Loveridge et al. 2006), combined

with models of potential ecological sustainability, the current rates of unsustainability

(Packer et al. 2009) are perhaps instead anthropogenically determined.

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3 Trophy  hunting  of  lions:  Conservation  threat  

3.1 Limitations  and  vulnerability    

Given current practices and unsustainable off takes, trophy hunting can also be a

significant threat to lion conservation. Trophy hunting can increase the vulnerability

of lions to other threats, which undermines the sustainability of the practice. Despite

models of projected resilience (Whitman et al., 2007) and the social ‘safety net’ that

prides can provide (Packer et al. 2009), in practice, reducing the population (even if it

is temporarily ‘sustainable’), nonetheless increases their vulnerability to a variety of

threats. Several authors also note the danger of environmental-based vulnerability.

Smaller and isolated lion populations are more vulnerable to disease (Packer et al.,

2011). The increased game fences surrounding hunting areas (often to prevent prey

species predation from wild dogs and cheetahs [Lindsey et al., 2005]) restrict

movements of wild populations and contain individual or multiple prides within

smaller discrete biogeographical units; this increases the risk of a synergistic interplay

of detrimental drivers (Lindsey et al., 2007). Croes et al., (2011) recently found that

in Cameroon, hunting zones outside of Pas act as population ‘sinks’, drawing lions

from the protected ‘source’ population.

Moreover, one (human-induced) form of vulnerability could be the relatively untested

‘anthropogenic Allee effect’ (Courchamp et al., 2006), whereby the value of the

charismatic lion will increase its rarity, which could ultimately lead to an extinction

vortex (ibid; Palazy et al., 2011, 2012). Packer et al. (2009) highlight this risk using

examples in Zambia, Tanzania and Zimbabwe of the maintenance of off-takes

through compensatory increases in hunting effort, where ecological criteria such as

minimum age are broken. Finally, local communities can undermine the vulnerable

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and brittle sustainability of trophy hunting. Lichtenfeld (2005) suggests that the

interplay of trophy and Maasia-induced hunting and the loss of habitat and reductions

in prey populations, are probably the reason for the lower lion density in Loibor

Serrit, Tanzania. Croes et al. (2011) suggest that high poaching and harvest rates of

17.5% in the Bénoué Complex, lion populations are well below carrying capacity.

They propose a fiver-year hunting moratorium, which, although criticized for lacking

sufficient evidence (Joppa and Hutton, 2012), is further defended by de Longh (2012),

given the functional extinction of lions in the neighbouring Gashaka-Gumti complex

in Nigeria (Henschel et al. 2010). To assess the viability of trophy hunting as a

conservation tool, trophy hunting must be evaluated in the light of the broader threats

to lions (see Bauer et al., 2008), and how trophy hunting’s interaction with these

threats may synergistically affect the delicate and vulnerable balance of sustainability.

The conservation benefits of trophy hunting are further limited by the manner in

which it is practiced. An important array of conditions needs to be met to achieve

ecological sustainability, and these currently seem a long way off from universal

adherence. Loveridge et al. (2006: 236) call for an “institutional structure that is able

to implement regulations effectively”. This is crucial to ensure sustainable practices,

good governance, transparency, reduced corruption and that the economic benefits are

distributed fairly (Whitman et al., 2007). Successful levels of regulation and

enforcement are however unlikely, given the extent of privately-owned hunting land

(Loveridge et al., 2006), and the varying scales of self-interest and desire for profit or

trophies. Trophy hunting is an archetypal conservation issue, where a lack of

resources and competing interests (such as between the profit-making industry and

international conservationists; between lions and farmers’ livestock), hinder

enforcement, ecological monitoring, and equal benefit distribution (ibid).

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3.2 Deleterious  impacts    

There is good evidence that trophy hunting can reduce the rate of increase of

populations of lions, having deleterious impacts, and in this sense, is ultimately

unsustainable. The harvesting of 2.4% of the total African lion population (600 out of

around 25,000) “is not sustainable” (IUCN, 2009:12); trophy hunting therefore

remains a threat to lion conservation. The majority of studies suggesting a detrimental

impact on lions from trophy hunting, focus on Tanzania (e.g. Lindsey et al., 2006,

2007; Loveridge et al., 2006; Packer et al. 2009, 2011), which holds most of the

remaining large lion populations (Packer et al. 2011), and where a large proportion of

land allows some degree of hunting (Fig. 7). Between 1996 and 2008, lion harvests

have declined by 50% across Tanzania, thereby suggesting that total lion populations

have also decreased (Packer et al., 2011). Hunting areas with the highest initial

harvests suffered the steepest declines, indicates a correlation between trophy hunting

and population decrease (Fig. 8). Trophy hunting appears to have been the primary

driver of a decline in lion abundance in the Tanzania’s trophy-hunting areas (ibid).

Elsewhere, in the Northwestern Matetsi Safari Area of Zimbabwe, between 1974-

2004 male lion harvests also reached “exceptionally high” unsustainable levels of up

to 11 lions per 1000km2 (Packer et al., 2006:2) (Fig. 9).

This population decrease from trophy hunting is related to two problems in lion

trophy hunting. Firstly, there is a significant sex bias, with the majority of lions

harvested being males (Packer et al., 2011; Lichtenfeld, 2005; Grobbelaar and

Masulani, 2003). This sexually selective force has significant impacts on the

sustainability of trophy hunting (Whitman et al., 2007). Surrounding Zimbabwe’s

Hwange National Park, Loveridge and Macdonald (2001) found that 67% of mature

male lions were harvested from a study population covering 6000 km2 of the National

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Park, reducing the adult male proportion from 30% to 13%. Packer et al. (2011:151)

found this “unsustainable” proportion to be up to 28% year-1 in the late 1990s.

Secondly, shooting underage lions (below 5 years) has significant impacts. The

practice of baiting in Tanzania for example mainly attracts underage lions who have a

propensity to scavenge (Whitman, 2002); this increases the chance of younger lions

being shot, especially when there is pressure for the high-paying client to shoot at

least one trophy (Kiffner, 2008). Whitman (2007) model the wider demographic

effects of different aged males on females, showing a clear unsustainability when

lions below 5 years are shot (Fig.6).

Drawing the demographic significance of the gendered bias and underage shooting

bias together, it is clear that the social structure of lions plays an important role in

trophy hunting impacts. One major socially driven impact of trophy hunting

documented in lion populations is infanticide (Lichtenfeld, 2005; Loveridge et al.,

2007), which can have long-term impacts on multiple populations (Grinnell et al.,

1995; Whitman et al., 2004). Packer et al.’s (2009) comparison of simulation models

(and harvest data) between infanticidal lions and non-infanticidal North American

black bears, projects lion population declines from even moderate hunting, with no

such parallel for black bears. The paternal investment of lions means that a bias of

male harvests increases the rate of replacement (ibid). Male lions also play an

important protective role for prides. Cooper (1991) document an example of one

potential impact in Savuti, Botswana, where they found high levels of

kleptoparasitism by spotted hyaenas amongst female groups that were devoid of

(hunted) males. Thus, with the potential that killing just one individual from a social

species (such as lions) can affect not only a whole pride but its long-term viability

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(Tuyttens and Macdonald 2000), highlights the significance of even small scale

analysis, and the serious consequences of trophy hunting

Furthermore, the male bias in hunting selection can lead to males expanding their

home range (due to a lack of competition), which increases the potential for them to

move out of the reserve, and therefore become vulnerable to trophy hunting, as seen

in the Okavanga Delta, Botswana (Loveridge and Macdonald, 2001). It would be even

worse for lion populations to shoot an equal male-female ratio; however, the fact that

trophy hunting necessitates this selective imbalance, and that this imbalance can result

in deleterious impacts, highlights the vulnerability caused, and the likely

unsustainability of trophy hunting practice. Thus, ecologically speaking, trophy

hunting can have significant harmful effects on lions, which undermines the notion of

trophy hunting as a conservation tool.

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4 Conservation  Values     Modern conservation is about “people in nature” (Collins, 2004: 461), the ethos of

which is increasingly challenged and politicized. Trophy hunting is controversial, the

argument being polarized between nature-centered intrinsic values and

anthropogenic-based utilitarian values. Ecosystem services, for example, are valued

and determined in functional and financial terms. However, translating the accepted

commodification of carbon into the functional commodification of (nonhuman)

animal life is not so easily supported.

Ethical considerations are inescapable elements of conservation strategies- especially

trophy hunting (Loveridge et al., 2006). Animal welfare organisations (e.g. PETA)

argue that it is fundamentally wrong to kill living creatures- especially for sport.

Conservationists may instead take an ethological approach; they question the extent to

which hunted animals suffer (Jeppesen, 1987; Macdonald et al., 2000), the risk of

overhunting (Whitman, 2002) and whether hunting is morally compatible with

conservation. Unethical practices like baiting also negatively impact public

perception of trophy hunting as a conservation tool (Lindsey et al., 2007). Some

authors (e.g. Ehrenfeld, 1988) consider biodiversity as an intrinsic value and therefore

also a moral absolute in conservation. This absolute view however, is currently

limited in a milieu of wildlife consumption and messy tradeoffs (Polasky, 2008), and

therefore trophy hunting, like wider conservation practice, requires a politicized

process of tradeoffs.

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4.1 Intrinsic-­‐Instrumental  Trade  Offs  

Counter to Loveridge et al. (2006: 227), it is important to attend to “motivational

[ethical and intrinsic] issues” in conservation, which need to be placed in the context

of population/species/global conservation strategies, and used to aid increasingly

popular functional approaches. Indeed, trade offs are necessary in trophy hunting, and

serve to blend the polarized intrinsic-instrumental values in conservation and trophy

hunting. Interestingly, the intrinsic value of lions manifests in ostensibly polarized

ways through trophy hunting, and is felt strongly on both sides of the hunting debate

(Loveridge et al., 2006). Drawing on insights from Loveridge et al. (2006) and Jepson

and Whittaker (2002), those involved in the hunting industry (at least historically)

express(ed) a right to functionally benefit (experientially or economically) from the

lion’s death, whilst simultaneously desiring intellectual and aesthetic preservation and

assuming a moral obligation/conviction to ensure the species’ survival. This

destabilizes the intrinsic-instrumental binary, and raises questions of whether this

intrinsically based moral responsibility and limit depends on the right to benefit from

the wildlife ‘resource’, which thereby (in death) gains an instrumental value. Perhaps

then, intrinsic moral value depends on instrumental value (which paradoxically

derives from an intrinsic value)?

There is an increasing necessity to introduce “wider management systems needed to

deliver sustainable use and, if possible, incentive-driven conservation” (Hutton and

Leader-Williams, 2003: 215). Trophy hunting is a pragmatic tool in this respect. For

contemporary conservation, tradeoffs are a significant aspect of translating ecological

ideals into conservation practice (Leader-Williams et al., 2010). Whether it is

balancing the short-term economic benefits of trophy hunting with the long-term risk

of exploitation (Lavigne et al. 1996), accepting loss of individuals as the price paid

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for providing a sustainable population in an area where no lions would otherwise

exist, or balancing environmental conservation with the development needs of local

communities, the conservation role of trophy hunting must be evaluated relative to the

alternative. One example of this potential tradeoff process is problem animal control.

Lindsey et al. (2006) also found strong interest among hunting clients to shoot

problem animals (those threatening humans or livestock), with Conover (2002)

lauding the potential for this to minimize the costs of living with problem animals

through economic compensation of local communities living near hunting areas in

Zimbabwe (Whitman et al., 2007). Another example is green hunting, whereby

clients pay huge prices to immobilize wildlife (Loveridge et al., 2006). More

compromising solutions, such as these, are one way of attending to both sides of the

debate.

Trophy hunting is not an ideal conservation strategy, but is an important tool, and

therefore shouldn’t be universally banned. Indeed, this can be illustrated through a

comparison with contemporary critiques of aid (see Moyo, 2009). Just like aid, trophy

hunting can be harmful to its recipients due to poor governance, corruption and

dependence; however, if it were suddenly taken away it would have even worse

consequences for the recipients (lions or aid-dependents). As economically quantified

by Lindsey et al., (2012), the huge profits generated from lions, means that totally

prohibiting trophy hunting would have even worse impacts on the lions (and

dependent ecosystems and local communities) than continuing the practice or even

over-hunting. For example, lion populations have declined in Kenya, since its trophy

hunting ban in 1977 (Lindsey et al., 2006). Thus, a pragmatic approach that does

attend to intrinsic and instrumental values is crucial to ensure trophy hunting of lions

is sustainable and beneficial to both lions and people.

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5 Conclusion   Trophy hunting can and does conserve lions. It incentivizes the protection of habitat

and maintenance of sustainable populations in areas where there is no conservation

alternative. However, increased vulnerability to other threats, poor governance,

corruption, and unethical and unsustainable practices pervasively undermine its

success as a conservation strategy, and can result in deleterious impacts. Trophy

hunting is a hugely significant conservation tool, although perhaps not a good

conservation strategy.

With such huge sums of money entering the industry, it is vital that the development

needs of local communities are met in order to justify what in practice may often be a

tradeoff between killing individual lions and benefitting locals. More studies of where

money actually accrues are therefore vital (A. Loveridge, 2011). Trophy hunting

necessitates a trade off between intrinsic moral values and killing. Trade offs of this

kind are common in contemporary conservation strategies- especially in conservation

outside of PAs (Willis et al., 2012)- which often balance instrumental values with

political and practical necessity. For trophy hunting to realize its conservation

potential and cease being a significant threat to lions, requires better use of scientific

information (Lindsey et al., 2006; Packer et al., 2011; Whitman et al., 2007), better

regulation, and better control mechanisms. Motivating all sections involved in trophy

hunting is crucial to achieve this. There is thus still a long way to go before trophy

hunting becomes a tool, and not a threat, to lion conservation

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6 References   Adams, W.M. (2004) Against Extinction: the story of conservation, Earthscan,

London.

Bauer, H., and van der Merwe, S. (2004) Inventory of free-ranging lion Panthera leo

in Africa. Oryx, 38(1): 26–31.

Bauer, H., Nowell, K., Packer, C. (2008) Panthera leo. IUCN Red List of Threatened

Species (IUCN, 2008), Gland, Switzerland. Available from:

http://www.iucnredlist.org/apps/redlist/details/15951/0 [accessed online: 10th

November 2011]

Bélair C., Ichikawa K., Wong B.Y. L., and Mulongoy K.J. (2010). Sustainable use of

biological diversity in socio-ecological production landscapes. Background to the

‘Satoyama Initiative for the benefit of biodiversity and human well-being.’

Secretariat of the Convention on Biological Diversity, Montreal. Technical Series,

52.

Bond, I., Child, B., de la Harpe, D., Jones, B., Barnes, J. and Anderson, H. (2004)

Private land contribution to conservation in South Africa. Parks in transition, (ed.

by B. Child), pp. 29-61. UK: Earthscan

Booth, V. (2002). Analysis of wildlife markets (sport hunting and tourism). WWF-

SARPO report, Harare.

Caro, T., C. R. Young, A. E. Cauldwell, and D. D. E. Brown. 2009. Animal breeding

systems and big game hunting: models and application. Biological Conservation

142(4): 909–929.

Chardonnet, P. (2002) Conservation of the African Lion: Contribution to a Status

Survey. International Foundation for the Conservation of Wildlife, France and

Conservation Force, USA. Accessed online: 29/2/12. [Available from:

http://www.conservationforce.org/pdf/conservationoftheafricanlion.pdf]

Collins, M. (2004) Against Extinction: The Story of Conservation (ed. By W.

Adams). Oryx 38(4): 461-462

Conover, M. 2002. Resolving human-wildlife conflicts: the science of wildlife

management. CRC Press, Boca Raton, Florida. Convention on Biological Diversity

(2003). Accessed online 4/3/12. [Accessible from:

http://www.cbd.int/doc/meetings/sbstta/sbstta-09/information/sbstta-09-inf-06-

en.pdf]

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Cooper, S.M. (1991) Optimal hunting group size: the need for lions to defend their

kills against loss to spotted hyaenas. African Journal of Ecology, 29(2): 130-136

Courchamp F, Angulo E, Rivalan P, Hall RJ, Signoret L, et al. (2006) Rarity Value

and Species Extinction: The Anthropogenic Allee Effect. PLoS Biology 4(12): e415

Croes, B., Funston, P., Rasmussen, G., Buij, R., Saleh, A., Tumenta, P.N. and de

Iongh, H.H. (2011) The impact of trophy hunting on lions (Panthera leo) and other

large carnivores in the Bénoué Complex, northern Cameroon. Biological

Conservation, 144(12): 3064-3072

Damm, G.R. (2008) Recreational Trophy Hunting: What do we know and what

should we do? Best Practices in Sustainable Hunting- A guide to best practices

from around the world. CCIC: International Council for Game and Wildlife

Conservation (ed. by R.D .Baldus, G.R. Damm, and K. Wollscheid), pp. 5-11.

[Accessible from: http://www.cic-

wildlife.org/fileadmin/Press/Technical_Series/EN/1_EN.pdf]

De Longh, H. (2012) The impact of trophy hunting on lions (Panthera leo) and other

large carnivores in the Bénoué Complex, northern Cameroon. Biological

Conservation, in press. Accessed online: 9/4/12. [Accessible from:

http://dx.doi.org/10.1016/j.biocon.2012.02.012]

Ehrenfeld, D. 1988. “Why Put a Value on Biodiversity?” Biodiversity (ed. by E.O.

Wilson and F.M. Peter), 212-216. Washington, DC: National Academy Press.

Fjeldsa, J., Burgess, N.D., Blyth, S. and de Klerk, H.M. (2004). Where are the major

gaps in the reserve network for Africa’s mammals? Oryx, 38, 17–25.

Fryxell, J. M., Packer, C. McCann, K., Solberg, E.J. and Sæther, B.E. (2010) Harvest

cycles and the sustainability of wildlife populations. Science, 328: 903–906.

Funtowicz, S. and Ravetz, J. (1992) The good, the true and the post-modern. Futures,

24(10): 963-976

Grinnell, J., Packer, C. and Pusey, A. E. (1995) Cooperation in male lions: kinship,

reciprocity or mutualism? Animal Behaviour, 49(1), 95–105 .

Grobbelaar, C. and R. Masulani (2003). Review of offtake quotas, trophy quality and

'catch effort' across the four main wildlife species elephant, buffalo, lion and

leopard. WWF SARPO.

Grunblatt, J., Said, M., Wargute, P. and Kifugo, S.C. (1995) Department of resource

surveys and remote sensing aerial surveys databases: methods and products.

21

Ministry of Planning and National Development, Department of Resource Surveys

and Remote Sensing, Nairobi, Kenya.

Henschel, P., Azani, D., Burton, C., Malanda, G., Saidu, Y., Sam, M., Hunter, L.,

(2010) Lion status updates from five range countries in West and Central Africa.

Cat News, 52(1), 34–39.

Hutton, J.M. and Leader-Williams, N. (2003) Sustainable use and incentive-driven

conservation: realigning human and conservation interests. Oryx, 37(2): 215-226

IUCN (2009) La grande chasse en Afrique de l’Ouest: quelle contribution à la

conservation? (Big Game Hunting in West Africa. What is its contribution to

conservation?) PAPACO Studies Number 2

Jenkins, R.K.B., Maliti, H.T. and Corti, G.R. (2003) Conservation of the puku

antelope in the Kilombero Valley Tanzania. Biodiversity Conservation, 12(4), 787–

797.

Jeppesen, J.L. (1987) The disturbing effects of orienteering and hunting on roe deer

(Capreolus capreolus) at Kaloe. Danish Review of Game Biology 13(1): 1–24

Jepson, P. and Whittaker, R.J. (2002) Histories of protected areas: internationalisation

of conservationist values in their adoption in the Netherlands Indies. Environment

and History, 8(2): 129-172.

Jew, E. and Bonnington, C. (2011) Socio-demographic factors influence the attitudes

of local residents towards trophy hunting activities in the Kilombero Valley,

Tanzania. African Journal of Ecology, 49(3): 277–285

Joppa, L. and Hutton, J. (2012) Trophy hunting of lions in West Africa. Biological

Conservation, in Press. Accessed online: 9/4/12. [Accessible from:

http://dx.doi.org/10.1016/j.biocon.2012.02.010]

Kiffner, C. (2008) African lions and the trophy hunting dilemma. Endangered Species

Update, 25(1), 3-8

Laing, A. (2012) Donald Trump's sons under fire over Africa hunting trip. The

Telegraph. Accessed online: 14/3/12. [Accessible from:

http://www.telegraph.co.uk/news/worldnews/africaandindianocean/zimbabwe/9143

550/Donald-Trumps-sons-under-fire-over-Africa-hunting-trip.html]

Lavigne, D. M., C. J. Callaghan, and R. J. Smith. 1996. Sustainable utilization: the

lessons of history. The exploitation of mammal populations (ed. by V.J. Taylor and

N. Dunstone), pp. 250-265. Chapman and Hall, London.

22

Leader-Williams, N. and Hutton, J.M. (2005) Does extractive use provide

opportunities to offset conflicts between people and wildlife? People and Wildlife:

conflict or co-existence? (ed. by R. Woodroffe, S.J. Thirgood, and A. Rabinowitz),

pp. 140-161. Cambridge: Cambridge University Press.

Leader-Williams, N., Adams, W.M. and Smith, R.J. (2010) Trade-offs in

Conservation: deciding what to save. Oxford: Wiley-Blackwell.

Leader-Williams, N., Milledge, S., Adcock, K., Brooks, M., Conway, A., Knight, M.,

Mainka, S., Martin, E.B. and Teferi, T. (2005) Trophy hunting of black rhino

Diceros bicornis: proposals to ensure its future sustainability. Journal of

International Wildlife Law and Policy, 8(1): 1-11

Lewis, D. M., and P. Alpert. 1997. Trophy hunting and wildlife conservation in

Zambia. Conservation Biology 11(1): 59–68.

Lichtenfeld, L.L (2005) Our Shared Kingdom At Risk: Human – Lion Relationships in

the 21st Century. PhD Thesis, Yale University.

Lindsey, P. (2008) Trophy hunting in sub-Saharan Africa: economic scale and

conservation significance. Best Practices in Sustainable Hunting, 11(1): 41-47

Lindsey, P.A., du Toit, J.T. and Mills, M.G. (2005). Attitudes of ranchers towards

African wild dogs Lycaon pictus: conservation implications for wild dogs on

private land. Biological Conservation, 125(1): 113–121.

Lindsey, P.A., Roulet, P.A., and Romañach, S.S. (2007) Economic and conservation

significance of the trophy hunting industry in sub-Saharan Africa. Biological

Conservation, 134(4): 455-469

Lindsey PA , Balme GA , Booth VR , Midlane N (2012) The Significance of African

Lions for the Financial Viability of Trophy Hunting and the Maintenance of Wild

Land. PLoS ONE 7(1): e29332

Lindsey, P.A., Alexander, R., Frank, L.G., Mathieson, A. and Romañach, S.S. (2006),

Potential of trophy hunting to create incentives for wildlife conservation in Africa

where alternative wildlife-based land uses may not be viable. Animal Conservation,

9(3): 283–291

LionAid (2012) Update on the LionAid Conference on the conservation needs and

status of African lions. Accessed online: 5/4/12. [Accessible from:

http://www.lionaid.org/campaign/2012/02/lionaid-to-host-a-conference-on-the-

conservation-needs-and-status-of-african-lions.htm]

Loveridge, A. (2011) Email. 9th March, 2011.

23

Loveridge, A. and Macdonald, D. (2001) Impact of trophy hunting in Zimbabwe. In

Lion Conservation Research. Workshop 2: Modelling Conflict (ed. by A.J.

Loveridge, T. Lynam and D.W. Macdonald), pp. 18– 19. October, Okavango Delta,

Botswana.

Loveridge, A.J., Reynolds, J.C., Milner-Gulland, E.J., (2006). Is sport hunting part of

conservation? Key Topics in Conservation (ed. by Macdonald, D.W., and Service,

K.), pp. 224-240.

Loveridge, A.J., Searle, A.W., Murindagomo, F., Macdonald, D.W. 2007 The Impact

of Sport-Hunting on the Population Dynamics of an African Lion Population in a

Protected Area. Biological Conservation, 134(4): 548-558

Macdonald, D.W., Tattersall, F.H., Johnson, P.J., et al. (2000) Management and

Control of Populations of Foxes, Deer, Hares, and Mink in England and Wales,

and the Impact of Hunting with Dogs. Published with Home Office (2000). The

Stationery Office, London. [Available online from:

http://www.huntinginquiry.gov.uk/mainsections/research/macdonaldfinal.pdf]

Milner-Gulland, E.J., Bukreeva, O.M., Coulson, T., Lushchekina, A.A., Kholodova,

M.V., Bekenov, A.B. and Grachev, I.A. (2003) Reproductive collapse in saiga

antelope harems. Nature 422(6928): 135

Moyo, D. (2009) Dead Aid: Why Aid Is Not Working and How There Is a Better way

for Africa. New York: Farrar, Straus and Giroux.

Myers, N. (1975) The silent savannahs. International Wildlife, 5(5): 5-10.

Nowell, K., and Jackson, P. (1996) Wild Cat: Status Survey and Conservation Action

Plan. IUCN Publications, Cambridge, UK.

Packer, C., Whitman, K., Loveridge, A., Jackson, J., Funston, P. (2006) Impacts of

trophy hunting on lions in East and Southern Africa: Recent off take and future

recommendations. Background paper for the Eastern and Southern African Lion

conservation workshop. Johannesburg, South Africa, pp. 1-15. 11-13th January,

2006.

Packer, C., Brink, H., Kissui, B., Maliti, M., Kushnir, H. and Caro, T. (2011) Effects

of Trophy Hunting on Lion and Leopard Populations in Tanzania. Conservation

Biology, 25(1): 142–153.

Packer, C., Kosmala, M., Cooley, H.S., Brink, H., Pintea, L., Garshelis, D., Purchase,

G., Strauss, M., Swanson, A., Balme, G., Hunter, L. and Nowell, K. (2009) Sport

24

Hunting, Predator Control and Conservation of Large Carnivores. PLoS ONE 4(6):

e5941.

Palazy L, Bonenfant C, Gaillard J-M, Courchamp F (2011) Cat Dilemma: Too

Protected To Escape Trophy Hunting? PLoS ONE 6(7): e22424

Palazy, L., Bonenfant, C., Gaillard, J.M and Courcham, F. (2012) Response: rarity,

trophy hunting and ungulates. Animal Conservation, 15(1): 16–17

People for the Ethical Treatment of Animals (2012) Why sport hunting is cruel and

unnecessary. Accessed online: 5/3/12. [Accessible from:

http://www.peta.org/issues/Wildlife/why-sport-hunting-is-cruel-and-

unnecessary.aspx]

Polasky, S. (2008) Valuing Nature: Biophysical or Monetary Measures? Economics

and Conservation in the Tropics: A Strategic Dialogue, January 31 – February 1,

2008

Ray, J.C., Hunter, L.T.B., Zigouris, J., (2005). Setting Conservation and Research

Priorities for Larger African Carnivores. Working Paper 24. Wildlife Conservation

Society, New York.

Tuyttens, F.A.M. and Macdonald, D.W. (2000) Consequences of social perturbation

for wildlife management and conservation. Behaviour and Conservation, (ed. by

L.M. Gosling and S.W.J. Cambridge), pp. 315–329. Cambridge University Press,

Cambridge.

Van Rijssel, J. (2008) Lion (Panthera leo) ecology and livestock conflicts in Waza

National Park, Cameroon. Environment and Development Sutdent Report no. 233.

Whitman, K. L. 2002. Safari Hunting of Lions: A review of policies, practices and

industry concerns. Proceedings of the 2nd meeting of the African Lion Working

Group, 2002 (ed. By H.O. De Waal), pp. 111-124. World Conservation Union,

African Lion Working Group, Brandhof, Republic of South Africa.

Whitman, K., Starfield, A.M., Quadling, H.S. and Packer, C. (2007) Modeling the

Effects of Trophy Selection and Environmental Disturbance on a Simulated

Population of African Lions. Conservation Biology, 21(3): 591–601

Whitman, K., Starfield, A.M., Henley, S., Quadling, H.S. and Packer, C. (2004)

Sustainable trophy hunting of African lions. Nature 428(6979): 175-178

Wilkie, D.S. and Carpenter, J.F. (1999a). The potential role of safari hunting as a

source of revenue for protected areas in the Congo Basin. Oryx 33(4), 339–345.

25

Wilkie, D.S. and Carpenter, J.F. (1999b). Can nature tourism help finance protected

areas in the Congo Basin? Oryx 33(4), 332–338.

Willis, K.J., Jeffers, E.S., Tovara,C., Long, P.R., Caithnessc, N., Smitd, M.G.D.,

Hagemann, R., Collin-Hansen, C. and Weissenberger, J. (2012) Determining the

ecological value of landscapes beyond protected areas. Biological Conservation,

147(1): 3-12

7 Figures  

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Figure 2: Proportion of hunters visiting each country (based on latest estimates for hunters visiting each nation). Source: Lindsey et al. (2007)

Figure 1: Recent trends in the number of foreign hunters visiting southern and East Africa, demonstrating an overall increase in total foreign hunters between 19990 and 2004. Source: Lindsey et al. (2007)

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Figure 3: Effects of 30 years of lion trophy hunting as a function of hunting quota size and male age in a hypothetical population. Average outcome after 100 simulation runs is shown from shooting males in four age groups (≥ 3, ≥4, ≥5, and ≥ 6 years old): (a) number of adult females after 30 years, (b) total number of males harvested, (c) total number of 5- to 6-year-old “trophies” harvested. Figs. 3a-c demonstrate that irrespective of quota size, harvesting males at least 5 years old had a negligible effect on population viability. Figs. 3a,b demonstrates the higher vulnerability of females than males. Fig. 3c suggests that placing age restrictions on lions shot increased the total number of males harvested after 30 years and increased the number of 5- and 6- year old trophies in the population by protecting young males. Source: Whitman et al., (2004, 2007)

Figure 4: Age estimation for adult lions using nose colouration. a, Identification photograph of a 3-yr-old Serengeti male; b, Excised photo of nose tip; c, GIS rendering of nose colouration; d, Age-change of nose colouration for males and females in two separate populations. Whitman et al., (2004) demonstrate the clear relationship between nose pigmentation and lion age, with the model proving “robust…to errors in age assessment based on nose coloration”. There is an obvious difference in the degree of pigmentation between 30% and 60%, and even if hunters applied a 50% rule (for 5-year-old lions), the long-term effect on population size would be small (Whitman et al., 2007). Professional hunters in Tanzania are currently using the nose pigmentation index in concert with other aging methods to reduce the likelihood of shooting immature male lions (ibid). Source: Whitman et al., (2004)

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Figure 5: Effects of 40 years of trophy hunting of male lions on female population size as a function of hunting quota size and male age. Average outcomes over 100 simulation runs from shooting males ≥3, ≥4, ≥5, and ≥6 years old are shown. The dotted lines show use of nose coloration as an accurate and reliable measure of male age: ≥40% black, 4 years old; ≥50% black, 5 years old; and ≥60% black, 6 years old. Source: Whitman et al. (2007)

Figure 6: Stochastic model using 40 years of data from the Serengeti. It shows the effects of 30 years of trophy hunting of male lions on the number of adult females as a function of the hunting quota size and male age for a population recovering from a hypothetical environmental disturbance after 100 simulation runs (males shot in four age groups: ≥3, ≥4, ≥5, and ≥6 years old). The graph demonstrates that provided age limits are followed, environmental disturbance did not undermine population viability/sustainability. Source: Whitman et al., (2007)

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Figure 7: Boundaries of different types of protected areas in Tanzania. Hunting is prohibited in National Parks, permitted in some parts of game reserves and permitted in ‘Game Control Areas’ and ‘Open Areas’. (Source: Pelkey et al., 2000). Trophy hunting is the chief land use in game reserves (aside from limited photographic tourism in some areas like the Selous Game Reserve) (Lichtenfeld, 2005). Tanzania has an extensive network of national parks (38,365 km2, including Ngorongoro Conservation Area), game reserves (102,049 km2), and game-controlled areas (202,959 km2), and has more lions than any other country in Africa. Source: Packer et al. (2011).

Figure 8: Average number of lions (heavy lines, diamonds) and leopards harvested (thin lines, circles) in major hunting areas. (The solid regression line demonstrates statistically significant declines between 1996 and 2008; dashed regression line demonstrates an insignificant relationship). Source: Packer et al. (2011)

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Figure 9: Estimated proportion of the lion population in each country removed annually by trophy hunting, using population estimates from Bauer and van der Merwe, 2004. Graph demonstrates that whilst this proportion of harvested lions is low for most countries (<4% per year- ibid), Zimbabwe seems to have harvested a far higher proportion of lions from 1988 to 2004 than any other country, and its off take rate has been 2-3 times higher than most other countries from 1998-2004. Source: Packer et al. (2009)