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

Review:

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

ABSTRACT

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

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.

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

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

4 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 (MilnerGulland 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, humanlion 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

5 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.

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

7 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

8 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 Tanzaniaâ&#x20AC;&#x2122;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 (LeaderWilliams and Hutton, 2005); there, hunting revenues dropped by only 12% (compared to the tourism industryâ&#x20AC;&#x2122;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 Â 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

9 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.

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

11 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).

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 19742004 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

13 Park, reducing the adult male proportion from 30% to 13%. Packer et al. (2011:151) found this â&#x20AC;&#x153;unsustainableâ&#x20AC;? 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.â&#x20AC;&#x2122;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

14 (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.

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.

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

17 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 shouldnâ&#x20AC;&#x2122;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.

5 Conclusion Â 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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Figures Â

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)

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

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)

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)

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)

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)