Biodiversity conservation seems to progress along a trajectory over time and economic development from gazetting isolated protected areas (often paper parks), to intensively managing those areas, to linking them via corridors, and finally to restoring those areas to some preconceived benchmark. Here, we describe this transition in South Africa and show how it now leads the world in largescale, biodiversity restoration, with lions (Panthera leo) at the forefront.

Although southern Africa is considered the cradle of humankind by many, it was not until a few thousand years ago that large scale transformation, through the use of fire and hunting, of the landscape begun. This was expedited a few hundred years ago when Europeans came to South Africa. They quickly changed the whole landscape with fenced farms, and created towns. With this they eradicated many of the large mammals; some such as the bluebuck (Hippotragus leucophaeus) were hunted to extinction. South Africa used to have huge migrations of springbok (Antidorcas marsupialis), creating some of the largest herds of mammals ever. The fences and hunting stopped this too, with the last recorded migration in 1896.

With the obvious dangers of large carnivores to humans and livestock, they were severely affected by this widespread increase in development. Along with the arrival of these farms over almost the entire landscape, many people, but particularly those in South Africa, lost the culture of living with dangerous animals, as is done in some other parts of Africa and India. Lion numbers were severely reduced to the point of extinction in South Africa, as they were in most parts of the world, with populations invariably restricted to a few small pockets. Some areas survived without too much development, perhaps only because of the high prevalence of diseases, such as malaria or sleeping sickness.

Around 1900, there was the proclamation of some of the oldest reserves in the world, such as the Kruger National Park and Hluhluwe-iMfolozi Park (HiP). HiP was originally a royal hunting ground for the Zulu kingdom, but was established as a park in 1895. The Kruger park area was first proclaimed in 1898 as the Sabie Game Reserve by the then president of the Transvaal Republic, Paul Kruger. In 1926, the National Parks Act was proclaimed and with it the merging of the Sabie and Shingwedzi Game Reserves into the Kruger National Park which, without the linked surrounding areas, is 2 million ha in size (20,000 sq. km). Since then, but especially after WWII, the number of protected areas in South Africa has grown considerably. These were mostly small (< 100,000 ha) but they began to restock them with the original wildlife, some including a few extralimital species or even exotics. Some extralimital species still remain but many of the exotics have been removed.

Some of these reintroduction exercises were massive in scale. Madikwe Game Reserve for instance reintroduced more than 8000 animals of 28 species between 1991 and 1997, one of the largest game translocation exercises in the world. This included lions.

The persecution of large carnivores, however, continued, even in conservation areas such as the Kruger National Park. Conservation staff would shoot African wild dogs Lycaon pictus on sight, and even help farmers to kill them outside protected areas. The aim was to protect the “game” or the ungulates, and the carnivores were simply seen as a threat to the game.

In the 1960s, one of the most famous reintroduction conservation success stories in the world began—the resurrection through translocation and reintroductions of the almost extinct white rhinos (Ceratotherium simum). Only a few remained, most of them in KwaZulu-Natal province. Driven by local conservationists such as the legendary Ian Player, rhinos were transported all over the country, including to the Kruger National Park and even to Zimbabwe and overseas. This turned the tide on their decline and until recently they seemed secure with numbers increasing to around 19,000. However, the last three years has seen a new onslaught of poaching and a new rhino war is been fought by rangers, especially in Kruger National Park. There are now again fears for their future and a renewed effort to transfer them to new places, again even outside South Africa, has begun. Black rhino (Diceros bicornis) similarly had their numbers decrease to about 2300 in 1993 but then they were similarly managed through the Black Rhino Expansion Programme which took their numbers up to about 5000. They are now also victims to the new wave of poaching and are potentially in trouble again.

For some of the large carnivores, a similar expansion exercise has been carried out. Much of the movement has not been purely for conservation purposes but rather to increase tourism potential of smaller reserves. Many tourists like to see the “Big Five” (lions, leopard (Panthera pardus), buffalo (Syncerus caffer), elephants (Loxodonta africana), and rhino). Similarly hunters in Africa like to say they have hunted in a Big Five area or even hunted the Big Five. For this buffalo, elephant, lion, white rhino (black rhino if just for tourism) and leopard are often introduced. Many of these translocations were done with little planning as to the potential future effects of these species in small fenced reserves. For instance, in many areas, the large carnivores quickly ate more than was sustainable and ungulate numbers needed to be supplemented at high costs. Similarly elephant numbers are approaching levels in some areas that will necessitate management so as to preserve other biodiversity in these reserves. Reserves have begun to move the elephants out or put them on contraceptives. The situation in many areas is not sustainable as moving elephants that have already been moved once sometimes causes behavioural problems. Also, there are not enough areas left needing elephants and moving them is very expensive. There are also obvious ethical considerations to the option of culling, which at some point may need to be considered.

In a more formal approach, similar in some ways to the coordinated rhino reintroductions and translocations, a plan to manage wild dogs was set up in 1997. An African Wild Dog PHVA (Population and Habitat Viability Analysis) was done which resulted in a plan for wild dogs to be reintroduced to a number of small reserves. These wild dogs were to be monitored intensively and management implemented to simulate a natural metapopulation. While the original aim of this exercise was to have 9 packs in the metapopulation, there are at present around 14 packs and 237 individuals. Each sub-population is managed by translocating individuals or groups to simulate natural pack dynamics and dispersals between subpopulations. Some individuals have left the subpopulations and now roam outside reserves, while some are in transit in this managed metapopulation. The only other population of wild dogs in South Africa is in Kruger National Park which is not managed and is currently estimated at 280 individuals including pups and those in the western boundary reserves.

Similarly, cheetahs have been managed between these small reserves to try and simulate natural processes. This is done by the National Cheetah Conservation Forum (NCCF). They aim to implement a co-ordinated relocation strategy that will: 1) ensure the long-term viability of cheetahs in small fenced reserves, 2) ensure the long term genetic and demographic integrity of the metapopulation, 3) increase the resident range of cheetahs in South Africa and 4) maximise the conservation benefits of cheetahs in small fenced reserves.

Lions were historically distributed across most of South Africa but were reduced to a few isolated areas by the early 1900s. Although presently under review, they are listed as Vulnerable on the IUCN Red List. In KwaZulu-Natal province, for instance, they became extinct in around 1938. Then, without human assistance, a male possibly migrating from Kruger turned up in iMfolozi in 1958. More were then reintroduced in March 1965. The population grew to about 140 animals in 1987 but then started declining and showed possible signs of inbreeding depression during the 1990s. To reverse this, 16 lions from Madikwe Game Reserve and Pilanesberg National Park, were introduced from 1999 to 2002. At the same time as this, lions were introduced into many other reserves around South Africa.

With the political changes in the early 1990s came a new wave of increased tourism to South Africa. This resulted in the increase in the number of reserves wanting lions on their property. Lions have now been reintroduced into over 40 small reserves in South Africa. The dilemma is that in small fenced reserves, the natural mechanisms of lion population limitations and predator-prey dynamics are absent. In open systems, ecological and behavioural processes limit population growth. Like elephants, many lion populations are now on contraceptives. Many reserves have removal programs to limit their numbers to maintain as natural a system as possible given the size of the reserves. In South Africa, lions are hunted for sport or lion bones. There is controversy around this as some situations are called canned-lion hunting where lions are specifically bred to be hunted, and are hunted in small areas. When considering lion conservation, we do not include these lions.

Lions when moved are kept in bomas at the site of release—a soft release, which improves survival. Post-release monitoring with the aid of radio or satellite collars is essential, especially in reserves with a high human density nearby. Often human activity is the cause of post-release mortality via direct persecution following human-wildlife conflict. In the end, however, what will affect the long-term persistence of lions in South Africa is a stable economy and governance. Factors affecting translocation success are removing the factors driving the initial population extinction and supportive social contexts. Lions were largely driven extinct in South Africa by direct persecution and loss of prey base. Direct persecution is understandable given the threat lions pose to human life and livelihood, and so conservation practices aim to ameliorate this by separating people from dangerous wildlife in South Africa through the legislated use of conservation fencing. The restoration of wildlife numbers has improved the status of the prey base for lions. The economic value of large wildlife ensures it is the general public/private sector largely driving the increase in lion numbers in South Africa today, particularly because wildlife is privately owned and individual entities can make money from owning them via regular game sales. There is wide-spread support for these actions.

South Africa has led the way in faunal restoration and this has led to the situation where the country’s protected areas cover 6% of the land, while an additional 13% is largely protected via the private sector as game ranches or ecotourism ventures. This is a boon for conservation. Most southern African countries have similar levels of private conservation land ownership. Key drivers for the success of conservation activities in this region is the willingness of conservationists to undertake drastic interventions, the support of government in these actions, and the economic benefits that have been seen from returning land to conservation from pastoralism. South Africa illustrates to the rest of the world the value of adequately costing wildlife into national economics and performing active and intensive conservation management.

Counter arguments to the South African conservation management philosophy are that intensive management is costly, may alter ecological interactions, may convey a sense of unnaturalness through heavily managed (manicured) wildlife areas, and benefit sharing. Intensive management is costly, but South Africans realise it is a price worth paying to conserve their natural heritage. Furthermore, the private sector is driving much of the expansion of conservation lands because it is a profitable land use, and often more profitable than the pastoralism that was historically practiced. The current philosophy of South African governmental conservation managers is to restore ecological interactions to those before humans vastly altered ecosystems and drove species extinct; however some private game ranches (particularly hunting reserves) stock extralimital species and unusual colour morphs of species because they garner higher prices from hunters. This has raised substantial concerns from the conservation community.

Notwithstanding the challenges of defining ‘natural’ on a continent that has been impacted by humans for as long as the species has existed, the management of South African reserves promotes wild-ness in a way that it is invariably part of the attraction of the sites. In fact, most reserves have humans fenced in to allow wildlife to roam freely beyond human habitation (tourist camps). Finally, sharing the benefits of protected areas to local communities is somewhat belatedly receiving the importance it deserves and local communities jointly run some reserves, but often benefit through employment, schools, education, and healthcare. The costs of living alongside protected areas in South Africa are largely mitigated by the use of fencing; however, it is increasingly being recognised that humans derive enormous physical and health benefits from living alongside protected areas and so this is an unmeasured benefit to local communities. Ultimately, the value of South Africa’s conservation management approach is illustrated by the constant or increasing population densities within their conservation areas, and the increase in the number of conservation areas. This stands in stark contrast to many other lion range states.

References:

Hunter LTB, Pretorius K, Carlisle LC, Rickelton M, Walker C, Slotow R & JD Skinner. 2007. Restoring lions Panthera leo to northern KwaZulu-Natal, South Africa: short-term biological and technical success but equivocal long-term conservation. Oryx, 41(2), 196–204. doi:10.1017/ S003060530700172X.

Miller SM, Bissett C, Burger A, Courtenay B, Dickerson T, Druce DJ, Ferreira S, Funston PJ, Hofmeyr D, Kilian PJ, Matthews W, Naylor S, Parker DM, Slotow R, Toft M & D Zimmermann. 2013. Management of reintroduced lions in small, fenced reserves in South Africa : an assessment and guidelines. South African Journal of Wildlife Research, 43, 138–154.

Slotow R & LTB Hunter. 2009. Reintroduction decisions taken at the incorrect social scale devalue their conservation contribution: African lion in South Africa. In: MW Hayward & MJ Somers. (eds). The Reintroduction of Top-order Predators. Wiley- Blackwell, Oxford.

Trinkel M, Ferguson N, Reid A, Reid C, Somers M, Turelli L, Graf J, Szykman M, Cooper D, Haverman P, Kastberger G, Packer C & R Slotow. 2008. Translocating lions into an inbred lion population in the Hluhluwe-iMfolozi Park, South Africa. Animal Conservation, 11(2), 138–143. doi:10.1111/ j.1469-1795.2008.00163.x

Matt W Hayward is a senior lecturer at the College of Natural Sciences, Bangor University, UK, M. hayward@bangor.ac.uk.

Michael J Somers is the Director of the Centre for Wildlife Management at the University of Pretoria. He is working on large carnivore (lions, cheetahs and wild dogs) and rhino reintroductions in the region. He has published widely on the subject including co-editing a Wiley-Blackwell book on the reintroduction of top-order predators with Matt Hayward, Michael.Somers@up.ac.za.

Illustration & Infographic: Kalyani Ganapathy

Photographs : Matt W Hayward

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