It may be the end of an era for the captive lion industry in South Africa – what does this mean for the lion bone trade?
The South African government made a landmark decision in May 2021 to end the country’s controversial captive lion industry. One reason given by Barbara Creecy, South Africa’s environment minister, for shutting down the captive industry was ‘the risk that trade in lion parts poses to stimulating poaching and illegal trade’, while the official government report highlighted the risk of the ‘laundering of poached parts’ into the legal market.1 The decision was also shaped by the accusations of severe neglect levelled by conservationists and animal welfare groups for years against the captive industry. The government’s assessment also found that the negative associations of captive lions were damaging to South African ecotourism.
Courtesy of Julian Rademeyer
Yet whether the legal bone trade has stimulated lion poaching, and whether bones from poached wild lions really have been ‘laundered’, are fiercely debated issues. So too is the question of whether ending the captive lion industry will put wild lion populations at greater risk of poaching, as international demand for lion bone will likely continue.
The rise and fall of the lion bone industry
There are estimated to be as many as 12 000 lions in captive facilities in South Africa. These lions are used for hunting,2 lion interactions and petting, as well as for commercial trade in lion parts, principally bones.3 Since the first permit to export lion skeletons from South Africa under CITES (the Convention on International Trade in Endangered Species of Wild Fauna and Flora) was issued in 2008, an industry has emerged in which intermediary lion bone traders buy skeletons from various breeding and hunting facilities and sell them to buyers in East and South East Asia, where they are sold as an alternative to (or marketed as) tiger bones for use in traditional medicine.4 An estimated 98% of lion bone exports from South Africa between 2008 and 2015 were to Laos and Vietnam – countries deeply implicated in the illegal wildlife trafficking of species such as rhino.5 There is also local demand in South Africa for lion parts that are used in traditional African medicine.
NOTE: The cited study below filtered out errors and discrepancies present in the CITES database through comparison and meticulous interrogation of the records.
SOURCE: Data up to 2017 on declared lion bone exports as reported in Vivienne Williams and Michael ‘t Sas-Rolfes, Born captive: A survey of the lion breeding, keeping and hunting industries in South Africa, PLOS One, 2019, https://doi.org/10.1371/journal.pone.0217409. From 2018 onwards, data from CITES export database.
NOTE: No export permits were issued in 2019 and 2020, so the cumulative number of legal exports was 0 for these years. Export numbers are for individual countries to which South Africa exported, and not necessarily the final destination.There is believed to be significant trade from transit countries to China.
SOURCE: Reported data on skeleton exports (2008–2015) from Vivienne Williams et al, A roaring trade? The legal trade in Panthera leo bones from Africa to East–Southeast Asia, PLOS One, 2017, https://doi.org/10.1371/journal.pone.0185996. Data on skeleton exports for following years, and on teeth and claw exports, from CITES trade database.
The May 2021 decision acts on the recommendation of the High-Level Panel of experts on wildlife conservation.6 Most of the panel recommended an immediate halt to captive lion breeding, use of captive lions in tourism and the trade in derivatives such as bones, arguing that the industry causes more economic damage than benefit.7
Yet the panel could not reach a unanimous position. Two minority positions of the panel were also published, which did not recognize that laundering of poached parts is a major risk to wild populations, and recommended different ways of monitoring and regulating the captive industry and bone exports.8
Have wild lion bones been ‘laundered’ through the south african market?
Some interest groups have voiced suspicions that the South African market provides a cover for laundered bones.9 Stephen Palos, chief executive of the Confederation of Hunting Associations of South Africa (CHASA), said that he has little doubt that laundering of bones has taken place, though this is more likely to be through volumes of captive-bred bones being understated in export shipments than poached lion bones being deliberately included.10
But in the view of many conservation scientists and researchers, these suspicions are not backed up by evidence. ‘I will state quite emphatically that both the risk of and evidence for “laundering” of lion body parts through South African legal channels since [a quota was imposed by a CITES ruling in 2016] is negligible to almost non-existent’, said Michael ‘t Sas-Rolfes, an economist who studies legal and illegal wildlife markets and sat on the High-Level Panel.11
Photo: Courtesy of Julian Rademeyer
Photo: Courtesy of Julian Rademeyer
The legal channel for exporting lion bones via CITES from South Africa included several monitoring checks, including DNA testing of lion bones and measuring of skeleton weights, to ensure that the correct individual skeletons were included in export shipments. A 2021 study reviewing the compliance of CITES lion bone exports with these monitoring systems found that there were few instances of suspected criminal activity.12 In the view of ‘t Sas-Rolfes, the review showed that the system was rigorous enough that ‘there really wasn’t much scope for using that system for any significant scale of illegal trade of wild-harvested lions or other big-cat products’.13
Other laboratory-based techniques such as mass spectrometry can be used to differentiate between wild and captive-bred lion bones, and could provide the key to finding out whether laundering has taken place.14 Yet according to David Newton, southern Africa director of the wildlife trade monitoring group TRAFFIC, these techniques remain in early stages of development and have not been widely used in lion bone exports. Now that legal exports are to end, in his view, it might never be known whether laundering really was widespread or not.15
Some experts also question whether laundering would make sense from an economic perspective. The quota, imposed from 2017, was substantially lower than the number of skeletons that would be available from the captive lion industry. Laundering poached bones from wild lions would therefore involve additional cost and risk.16 ‘One has to look at the motivation for that. Why would you launder wild bones as captive-bred bones, when lions from captive populations are so easy to access and captive bones are just perfectly acceptable for the end destination?,’ said Newton.17
However, some conservationists report that a parallel, illegal trade has emerged, which may circumvent the CITES monitoring system. According to Kerri Rademeyer, CEO of the Zambian non-profit organization Wildlife Crime Prevention, this parallel trade takes place in multiple forms including ‘lion cake’ – a preparation of boiled down and compressed lion bone for medicinal use, which could be more difficult to track than the bones themselves.18
NOTE: Conservation experts are divided in opinion on whether it is possible to link reported increases in African lion killings with the South African trade in lion parts.
SOURCE: Paul Funston, Kris Everatt and Karen Wood, Report on the connection between the legal lion bone and body parts trade in South Africa and a rise in the poaching of wild and captive lions, Panthera.
Recent trends in lion poaching: is there a link to the bone trade?
Some conservationists believe that the legal trade in lion parts is stimulating international demand and leading to an increase in lion poaching, particularly in countries bordering South Africa. A key piece of evidence in this debate is a 2019 study of lion killings in Limpopo National Park, Mozambique, which neighbours South Africa’s Kruger National Park. The study found that the targeted poaching of lions for body parts accounted for all 61% lion mortalities, between 2011 and 2018.19
‘It looked like it was coinciding with South Africa’s legal export of body parts, because of its geographic nature and the fact that it was right next door to South Africa’, said Kristoffer Everatt, project manager for the Lion Program at Panthera, the global wild cat conservation organization, and an author of the study. According to Everatt, perceptions shifted in the Limpopo area around 2013–2014, from lion bones being a comparatively ‘worthless’ commodity, to parts such as bones, teeth and claws suddenly being perceived as high value.20 Further analysis by Panthera has compared rates of poaching in Limpopo to the number of lion export permits issued per year in South Africa.21 ‘The patterns there just were so similar. I have to believe there’s a link’, said Paul Funston, Lion Program Senior Director at Panthera, though acknowledging that it is difficult to prove this link statistically.22
Illegal trade in lion parts: an emerging picture
Reports of lion poaching for parts, and seizures of parts such as bones, teeth and claws, have been on the rise in some parts of southern Africa (as shown on the timeline).23 According to Carlos Lopes Pereira, head of Law Enforcement and Anti-poaching at Mozambique’s National Administration of Conservation Areas, illegal demand for lion parts has risen in areas across Mozambique.
Yet the data suggesting that there is a trend towards poaching for body parts – of any type – is by no means uniform. ‘Colleagues and I used long-term mortality data from around Ruaha in Tanzania, and Hwange in Zimbabwe, and thankfully found no evidence indicating a trend towards the killing of lions for commercial body parts’, said Amy Dickman, director of the Ruaha Carnivore Project.24 However, she adds, trends in different lion ranges may be very different. Others working and researching in Tanzania also argued that there was, as yet, no evidence there to suggest a trend towards poaching for body parts for international trade.25
Others disagree that an increase in poaching can be linked to the South African legal trade in lion parts. Lopes Pereira, for example, argued that the increase in poaching for parts in Mozambique only began several years after the trade became established in South Africa, making it hard to establish a direct connection with the legal trade.26 Instead, parts from poached and poisoned lions from Mozambique are largely sold on to Asian traffickers or smuggled to neighbouring countries including Zimbabwe, Zambia and Malawi before being shipped to Asian destination markets.27
‘I personally disagree with the narrative … that the poaching in Limpopo National Park is or was driven to any significant degree by the demand for lion bones and especially that South Africa’s legal trade somehow had a causal role to play here’, said ‘t Sas-Rolfes. In his view, poaching in this particular area was driven by a number of factors, such as protest-related killings of lions by communities as a backlash to the militarized approach to conservation taken in Limpopo National Park,28 and the presence in the area of wildlife trafficking networks that had previously dealt in rhino horn who could also have begun trafficking in other wildlife such as pangolin and lion body parts. ‘It is interesting to note that around the same time that lion poaching picked up [in Limpopo National Park] there was a surge in illegal trade in jaguar fangs in Latin America … and this also coincided with the move of Asian wildlife jewellery and trinket trade from physical markets to online platforms, giving those markets for items such as big-cat tooth and claw products wider reach’, he said.29
Funston, of Panthera, however, sees demand for claws and teeth for ‘trinkets’ as part of the same picture. ‘People have so habitually referred to the trade out of South Africa as lion bone trade, I think they forget and ignore the very significant number of teeth and claws that are going out as well’, he said. ‘[These are] stimulating a market, perhaps a different market than the market that bones would be used for, but nevertheless a market. [It] would seem to be lucrative enough that illegal hunters around the sub-region have started killing lions and removing faces, teeth and claws et cetera, from the carcasses that they hunt.’30
Everatt also acknowledged that other factors could be at play in Limpopo, including the presence of established rhino trafficking networks, meaning that there was already an established culture of poaching and poaching-related corruption in the region. ‘The same individuals that were poaching elephant and rhino are now also poaching lion, and often on the same trips … because they could make a trip to Kruger and on the way they would snare something, lay some poison, go hunting for elephant and rhino and then on their way home, they would pick up some lion body parts’, he said.31 Lopes Pereira also agreed that there is a convergence of networks dealing in wildlife that is in demand in Asia, and that rhino horn, ivory, pangolin and leopard skin and claws have all been seized alongside lion products.32
Not only is interpreting the data on lion poaching a challenge, it is also a challenge to collect this data in the first place. ‘One of the complexities here is that killings of lions (and other wildlife) often involves mixed motivations – so a lion might be killed because of conflict or cultural prestige, but then body parts also used for local or international trade’, explains Dickman. Another issue is identifying when lions have been killed. ‘If poachers are actually going to process a carcass, to the point where it’s got [only] bones left, law enforcement et cetera aren’t going to find the carcass in the wild’, says Funston.33 This makes it difficult to assess the number of lions being killed for parts, beyond the few instances where poachers have been arrested in possession of bones. This in turn makes defining poaching – and therefore measuring whether it is on the rise – more complicated than people usually imagine.
The future of lion poaching and illegal trade
Many experts thought it possible that shutting down the captive lion industry could lead to increased poaching of wild lions in future. ‘I am very worried about the potential unintended consequences of this decision’, said Dickman. ’This ruling would permanently shut off the only legal supply of lion bone, and yet we know the demand is still there, and may be rising. That leads to the obvious risk that demand will increasingly be met through illegal and unregulated killing of wild lions, which could pose a major threat for those populations.’34
However, some viewed the potential benefits as outweighing the risks. ‘The current trajectory proves that practices within the captive lion industry are irresponsible, inhumane and unsustainable. The industry in itself is a risk to wild lion population conservation and broader biodiversity conservation efforts,’ said Pricilla Stiglingh, of the South African NSPCA.35 ‘In mitigating risks [to wild lions], one cannot allow the practices of one industry to be detrimental to the larger biodiversity and ecotourism sector of South Africa. This needs to be looked at in a holistic manner by including all the threats wild lions face including use of bones and body parts, indiscriminate killing and human wildlife conflict and habitat loss. The solution is for all stakeholders, communities and government to commit to the overall conservation of wild lions by increasing suitable habitat and thriving wild lion populations, instead of defending a commercialized captive lion industry.’
Funston agreed. ‘Yes, we might expect increased rates of poaching’, he said. ‘[But] to suggest in any way that we should perpetuate an industry because we’re scared of the knock-on consequences of taking action, that’s just weak in my view.’36
Some also argued that the legal trade has already driven a demand for illegal supplies of lion bones and derivatives, and that the industry should be ended. ‘The whole lion bone trade may never have become a legitimate trade until South Africa made it one,’ said Rademeyer.37 Others disagreed, arguing that since international demand is already a reality, a more practical approach going forward would be an adaptive management approach, involving phasing out captive lions but making use of existing stockpiles of products such as bone and, if necessary, phasing in other sources such as wild-managed lions.38
Some facilities and traders currently exporting lion bone legally may turn to alternative, more clandestine ways of exporting bones from captive lions. ‘In terms of the illegal side, that is pretty much a no-brainer how things are going to play out in future’, says one lion bone trader. ‘The government made their own bed so they have to lie in it’.39 Palos, of CHASA, agreed, saying there is little doubt that some lion bone traders will be approached by individuals looking for illegal exports of bone.
Photo: Courtesy of Julian Rademeyer
A 2019 study surveying lion breeding facilities found that 52% of respondents indicated they would adapt by seeking ‘alternative markets’ for lion bones if the export was legally restricted.40 ‘We left it up to the respondents to interpret what “alternative markets” meant – and I’m not certain that they all interpreted this to mean illegal [exports of bones]’, said Vivienne Williams, a leading expert on the lion bone trade.41
In Funston’s view, the business links and networks formed in the legal lion trade could form the basis of illegal trafficking networks in future. This would be similar to the situation whereby, for many years, it was legal to hunt white rhino horn in South Africa. Asian rhino traders commissioned hunts to export horns legally, and during that time created vast networks of contacts that then transitioned into illegal rhino horn trafficking once rhino horn exports were banned. ‘I’m absolutely convinced that in the lion game, that the farms [and] traders that are now involved are very familiar with each other. They’re very used to and comfortable with trading [lion] products legally. If a country bans it … it’s very likely that the same people will just continue those networks and utilize those networks’, he said.42
Known wildlife traffickers have also been historically involved in the lion bone trade. One major early buyer, for example, was Vixay Keosavang,43 who headed one of the world’s largest wildlife trafficking syndicates that transported wildlife products such as rhino horn and ivory to Asia.44 However, according to Williams, as the industry grew over time and more people entered the lion bone trade, fewer breeders and traders had direct connections to these known wildlife traffickers.45
To be implemented, the recommendations of the High-Level Panel still need to be brought into legislation, which can be a time-consuming process.46 Any future legislation is also expected to be challenged in the courts and fought out between those with commercial interests in the lion industry and animal rights groups.
The different views among conservation experts on the decision to end the industry, and the impact this could have on illegal trade in lion parts, demonstrates how complex this policy question is, as it must balance the sometimes-conflicting interests of countering criminal activity, conservation, animal rights and the private sector. While the South African government may be striving to create evidence-based policy, much of the evidence on lion bone trade still appears very much up for debate.
Notes
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South African Department for Forestry, Fisheries and the Environment, Statement by Minister Creecy: Release of report of high-level panel 2 May 2021, https://www.environment.gov.za/speeches/creecy_releaseofhlpreport_pretoria; The High-Level Panel of Experts for the review of policies, legislation and practices on matters of elephant, lion, leopard and rhinoceros management, breeding, hunting, trade and handling, High-Level Panel Report – for submission to the Minister of Environment, Forestry and Fisheries, 15 December 2020, https://www.environment.gov.za/sites/default/files/reports/2020-12-22_high-levelpanel_report.pdf. ↩
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‘Canned hunting’ can take place in a variety of forms, including hunting an animal in an enclosed space or when it is tranquilized. Overall, the practice involves ‘unfairly preventing the target animal from escaping the hunter, thereby eliminating “fair chase” and guaranteeing the hunter a trophy’, as reported in the 2018 Parliamentary colloquium on captive lion breeding for hunting and lion bone trade, Committee Report, 13 November 2018, https://cer.org.za/wp-content/uploads/2018/11/Final-PCEA-Captive-Lion-Breeding-Colloquium-Report-13112018.pdf. ↩
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The High-Level Panel of experts advising the government on wildlife issues including the captive lion industry cited an estimate from a 2016 study of 8 000–8 500 captive lions in South Africa. (Tourism Research in Economic Environs & Society (TREES), Economic Value of the South African Private Lion Sector, 26, 2016). However, other organizations estimate that the total is far higher. Lion welfare and anti-captive-industry campaign organization Blood Lions estimates that there could be 450 facilities holding up to 12 000 lions (see Elizabeth Claire Alberts, South Africa pulls the plug on controversial captive lion industry, Mongabay, 3 May 2021, https://news.mongabay.com/2021/05/south-africa-pulls-the-plug-on-controversial-captive-lion-industry/). That there are such varied estimates of how many captive lions are currently in South Africa suggests that managing the shutdown of the industry and monitoring how lion bones are disposed of (to ensure none are exported illegally) could be a major challenge. ↩
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Vivienne L Williams and Michael J ‘t Sas-Rolfes, Born captive: A survey of the lion breeding, keeping and hunting industries in South Africa, PLoS ONE, 14, 5, 2019, e0217409, https://doi.org/10.1371/journal.pone.0217409. ↩
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The 2018 Parliamentary Colloquium on Captive Lion Breeding for Hunting and Lion Bone Trade, Committe Report, 13 November 2018, https://cer.org.za/wp-content/uploads/2018/11/Final-PCEA-Captive-Lion-Breeding-Colloquium-Report-13112018.pdf. ↩
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The High Level Panel was formed following a hearing of a committee on environmental affairs in 2018, in which it was reported that South Africa’s captive lion breeding industry ‘did not contribute to conservation and was doing damage to South Africa’s conservation and tourism reputation’, and a decision was made to convene a High-Level Panel in October 2019 to assess the industry. South African Department for Forestry, Fisheries and the Environment, Statement by Minister Creecy: Release of report of high-level panel 2 May 2021, https://www.environment.gov.za/speeches/creecy_releaseofhlpreport_pretoria. ↩
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The High-Level Panel of Experts for the review of policies, legislation and practices on matters of elephant, lion, leopard and rhinoceros management, breeding, hunting, trade and handling, High-Level Panel Report – for submission to the Minister of Environment, Forestry and Fisheries, 15 December 2020, https://www.environment.gov.za/sites/default/files/reports/2020-12-22_high-levelpanel_report.pdf. ↩
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The first of these minority views argued that it was possible to view the legal supply of lion bones as meeting consumer demand ‘that might otherwise be met through illegal means, including increased poaching of wild lions’, and that the ‘risk of laundering was effectively reduced to negligible levels’ in the 2017 and 2018 quotas. It recommended that future annual quotas, including possible zero quotas, follow a structured decision-making approach to minimize illegal activity and impact on wild populations, The second minority view argued that canned hunting and other captive lion businesses continue with the government establishing more oversight over lion breeders and other measures to regulate the industry more rigorously. ↩
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Interview with Pricilla Stiglingh, policy, regulations, compliance and research officer at the NSPCA, 11 May 2021, by phone. ↩
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Interview with Stephen Palos, CEO of the Confederation of Hunting Associations of South Africa, 19 May 2021, by phone. ↩
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Email correspondence with Michael ‘t Sas-Rolfes, economist, 25 May 2021. ↩
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Vivienne Williams et al., Monitoring compliance of CITES lion bone exports from South Africa, PLoS One, 16, 4, 2021, e0249306. ↩
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Email correspondence with Michael ‘t Sas-Rolfes, economist, 25 May 2021. ↩
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Peter Coals et al., DART mass spectrometry as a potential tool for the differentiation of captive‐bred and wild lion bones, Biodiversity and Conservation, 30, 2021, 1 825–1 854, https://doi.org/10.1007/s10531-021-02170-2. ↩
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Interview with David Newton, TRAFFIC southern Africa director, 5 June 2021, via Zoom. ↩
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Email correspondence with Michael ‘t Sas-Rolfes, economist, 25 May 2021. ↩
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Interview with David Newton, TRAFFIC southern Africa director, 5 June 2021, via Zoom. ↩
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Email correspondence with Kerri Rademeyer, CEO of Wildlife Crime Prevention, 17 May 2021. ↩
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KT Everatt, R Kokes and C Lopez Pereira, Evidence of a further emerging threat to lion conservation; targeted poaching for body parts, Biodiversity and Conservation, 28, 2019, 4 099–4 114, https://doi.org/10.1007/s10531-019-01866-w. Body parts were also removed in cases where lions were killed in conflict with livestock, further suggesting that there is an illicit demand for body parts. The lion population in Limpopo has reportedly ‘collapsed’ over the past decade. Interview with Kristoffer Everatt, conservation scientist and practitioner at Panthera, 11 May 2021, via Zoom. ↩
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Interview with Kristoffer Everatt, conservation scientist and practitioner at Panthera, 11 May 2021, via Zoom. ↩
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Paul Funston, Kris Everatt and Karen Wood, Report on the connection between the legal lion bone and body parts trade in South Africa and a rise in the poaching of wild and captive lions. Shared directly by the authors, 25 May 2021. ↩
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Interview with Paul Funston, director of the Lion Program at Panthera, 11 May 2021, by phone. ↩
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Vivienne L Williams et al., Questionnaire survey of the pan-African trade in lion body parts, PLoS ONE, 12, 10, 2017, https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0187060. ↩
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Email correspondence with Amy Dickman, director of the Ruaha Carnivore Project, 14 May 2021. ↩
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Interview with Tim Davenport, country director for Wildlife Conservation Society in Tanzania, 20 May 2021, via Skype; email correspondence with Amy Dickman, director of the Ruaha Carnivore Project, 14 May 2021; interview with David Newton, TRAFFIC southern Africa director, 5 June 2021, via Zoom. ↩
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Email correspondence with Carlos Lopes Pereira, head of Law Enforcement and Anti-poaching at Mozambique’s National Administration of Conservation Areas, 7 May 2021. ↩
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Ibid. ↩
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Rebecca Witter, Why militarized conservation may be counter-productive: illegal wildlife hunting as defiance, Journal of Political Ecology, 28, 1, 2021, 175–192, https://journals.librarypublishing.arizona.edu/jpe/article/id/2357/. ↩
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Email correspondence with Michael ‘t Sas-Rolfes, economist, 25 May 2021. ↩
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Interview with Paul Funston, Director of the Lion Program at Panthera, 11 May 2021, by phone. ↩
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Interview with Kristoffer Everatt, conservation scientist and practitioner at Panthera, 11 May 2021, via Zoom. ↩
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Email correspondence with Carlos Lopes Pereira, head of Law Enforcement and Anti-poaching at Mozambique’s National Administration of Conservation Areas, 7 May 2021. ↩
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Interview with Paul Funston, director of the Lion Program at Panthera, 11 May 2021, by phone. ↩
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Email correspondence with Amy Dickman, director of the Ruaha Carnivore Project, 14 May 2021. ↩
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Interview with Pricilla Stiglingh, policy, regulations, compliance, and research officer at the NSPCA, 11 May 2021, by phone. ↩
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Interview with Paul Funston, director of the Lion Program at Panthera, 11 May 2021, by phone. ↩
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Email correspondence with Kerri Rademeyer, CEO of Wildlife Crime Prevention, 17 May 2021. ↩
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Email correspondence with Michael ‘t Sas-Rolfes, economist, 25 May 2021. ↩
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Lion bone trader, speaking anonymously via email, 25 May 2021. ↩
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Vivienne L Williams and Michael J ‘t Sas-Rolfes, Born captive: A survey of the lion breeding, keeping and hunting industries in South Africa, PLoS ONE, 14, 5, 2019, e0217409, https://doi.org/10.1371/journal.pone.0217409. ↩
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Email correspondence with Vivienne Williams, researcher at the School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, 25 May 2021. ↩
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Interview with Paul Funston, director of the Lion Program at Panthera, 11 May 2021, by phone. ↩
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Vivienne Williams et al., Bones of Contention: South African trade in African Lion bones and other body parts, TRAFFIC, July 2015, https://www.traffic.org/publications/reports/bones-of-contention-south-african-trade-in-african-lion-bones-and-other-body-parts/. ↩
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Julian Rademeyer, Untouchable? Wildlife crime kingpin Vixay Keosavang, Environmental Investigation Agency, 14 February 2014, https://eia-international.org/blog/vixay-keosavang-an-untouchable-kingpin-of-wildlife-crime/. ↩
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Interview with Vivienne Williams, researcher at the School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, 26 May 2021, by phone. ↩
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Ibid. ↩