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Comparing South African and Canadian Practices for Managing Large Herbivores in Fenced Protected Areas

As human civilization expands and depletes resources, protected areas, such as national parks, have become key areas of conservation. Increasingly, protected areas are becoming ‘islands’ of wilderness.1,2,3 As a result, wildlife cannot move freely throughout their ranges, concentrating their impacts.20 Large herbivores greatly impact vegetation in areas they occupy;1,3,4,5,6 such impacts are compounded in fenced protected areas,1 especially i f predation is reduced or absent and if populations become hyperabundant.7 Therefore, wildlife managers must take a more active role. Kruger National Park (KNP) in South Africa and Elk Island National Park (EINP) in Canada are both fenced parks with large numbers of the largest herbivore species on their respective continents. KNP has the African elephant Loxodonta africana, listed as vulnerable,8 while EINP has the bison Bison bison. EINP has two subspecies of bison, wood bison B. b. athabascae and plains bison B. b. bison,9 both of which have threatened status in Canada.10 When too large or too small, the populations of these species could negatively impact overall biodiversity as it connects to ecosystem health and sustainability. There is limited literature on the management of highly abundant large herbivores in fenced protected areas on sites beyond Africa, and on comparisons between practices in Africa and North America. KNP and EINP offer an opportunity for comparison because of their similar management objectives, fencing policies, and issues with high profile populations of large herbivores that have large impacts on ecosystems. Although the parks are unique in some respects (i.e., different ecosystems, continents, and amounts of visitation), the fundamental aspects of wildlife management for large herbivores can be applied to other parks and countries. The purpose of this article is to compare past and present management strategies in KNP and EINP and the methods they employ with a goal of generating insights for improved management strategies.

As human civilization expands and depletes resources, protected areas, such as national parks, have become key areas of conservation. Increasingly, protected areas are becoming 'islands' of wilderness. 1,2,3 As a result, wildlife cannot move freely throughout their ranges, concentrating their impacts. 20 Large herbivores greatly impact vegetation in areas they occupy; 1,3,4,5,6 such impacts are compounded in fenced protected areas, 1 especially if predation is reduced or absent and if populations become hyperabundant. 7 Therefore, wildlife managers must take a more active role. Kruger National Park (KNP) in South Africa and Elk Island National Park (EINP) in Canada are both fenced parks with large numbers of the largest herbivore species on their respective continents. KNP has the African elephant Loxodonta africana, listed as vulnerable, 8 while EINP has the bison Bison bison. EINP has two subspecies of bison, wood bison B. b. athabascae and plains bison B. b. bison, 9 both of which have threatened status in Canada. 10 When too large or too small, the populations of these species could negatively impact overall biodiversity as it connects to ecosystem health and sustainability. There is limited literature on the management of highly abundant large herbivores in fenced protected areas on sites beyond Africa, and on comparisons between practices in Africa and North America. KNP and EINP offer an opportunity for comparison because of their similar management objectives, fencing policies, and issues with high profile populations of large herbivores that have large impacts on ecosystems. Although the parks are unique in some respects (i.e., different ecosystems, continents, and amounts of visitation), the fundamental aspects of wildlife management for large herbivores can be applied to other parks and countries. The purpose of this article is to compare past and present management strategies in KNP and EINP and the methods they employ with a goal of generating insights for improved management strategies.

Study Areas
KNP and EINP share common goals and histories (Table 1)  share some similar characteristics in regard to their impacts on ecosystems and they are both keystone species (Table 2). Nevertheless, the parks have different sizes and densities of their largest herbivores (Table  1). We recognize that it is difficult to select two parks with enough similar characteristics to allow for fair comparisons of their management strategies; we chose these parks because they are home to their continent's largest herbivores, the parks are fenced, and we have some familiarity with them.
Depending on definitions used, t h e s e t w o s p e c i e s m i g h t b e overabundant in their respective parks. Following Parks Canada's policy, a hyperabundant wildlife population meets one of the following conditions: 1) exceeds the upper range of natural variability and impacts ecological integrity, 2) has resulted from alteration of natural mechanisms for population regulation, 3) there is a risk of negatively affecting native species at risk, and 4) the population has caused impacts beyond the historical range of variation. 14 By this definition, bison in EINP are hyperabundant. Parks Canada is required to manage hyperabundant populations to maintain or restore ecological integrity. Methods to manage hyperabundant populations should best contribute to restoring or maintaining ecological integrity, mimic natural processes, and have been found to be effective on similar species. KNP also adheres to these standards and, though the term hyperabundant is not used in South African legislation, their National Norms and Standards for the Management of Elephants in South Africa 15 does outline the impacts elephants can have and what measures can be taken when management plan objectives are not met due to population size or distribution. KNP's elephant population is recorded to have impacts on vegetation and, therefore, other species 12 thereby hindering KNP's biodiversity management objective. 16 Therefore, given the impacts of the elephant population in KNP coupled with Parks Canada's definition and the information from the Norms and Standards 15 , both populations can be defined as hyperabundant.
The main methods employed to deal with abundant populations of large herbivores are translocation, contraception, culling, and nonmanagement. 1,3,13 Another less common method is distribution control. Non-management is not an option for elephants in KNP because their population is too large and is not an option for EINP because of its small land area and lack of predators. The remaining methods will be discussed in relation to the management of elephants in KNP and bison in EINP and how they are employed from different strategic points of view (i.e. impact control for KNP versus numbers control for EINP). When developing and implementing a wildlife management plan, park managers need to consider the advantages and disadvantages of each of these methods, as well as the behaviour of the species on which they are being applied.

Analysis of Management Methods
First, translocation is a nonlethal method of removing part of a population from a protected area while maintaining the species' genetic pool and re-establishing or increasing population numbers in a new area. Translocation can be difficult because it is expensive, requires a large and skilled team, is time-consuming, and can be difficult to find suitable and accessible translocation sites. 13,17 Elephants and bison are both large animals that require special handling and equipment to ensure the safety of animals and humans. Unlike bison,   12,17 This process led to the development of delinquent juveniles (ie. without the mentoring of adults) until translocations involving entire cow/calf herds and adult males was possible.
Translocation operations occur in cooler weather (Table 2), the animals are kept standing during transport, and arrive at new site in the daylight to reduce stress and injury. 17,18,19 Currently, there are few translocation areas available in Africa for elephants, because most areas have populations already, are not accessible, or are in politically unstable countries. However, KNP has effectively increased its size by opening up its fenced borders to neighboring game reserves, which then become contract national parks, and to the Mozambique's Limpopo National Park to form the Great Limpopo Transfrontier Park and Conservation Area. 16 Consistent management will require considerable inter-jurisdictional cooperation. EINP is a major source herd for bison re-introduction programs in North America and Europe because of the genetic purity and disease free status of the bison. 20 Second, contraception involves preventing the union of sperm and egg during sexual activity. The advantages are that no animals are killed, and it can be non-invasive and reversible. 3 Third, culling is an effective but controversial method. It quickly r e d u c e s l a r g e p o p u l a t i o n s , effectively controls disease, and can result in revenue from product sales. 1,3,13 However, culling causes a loss to the genetic pool, appears counterproductive when the broader population in question is listed as vulnerable or threatened 6,13 and can result in disturbing natural fluctuations in population numbers. 28 The most humane way to kill an elephant is with a brain shot, but this requires a skilled shooter due to the skull and muscle structure of the head. 12,13 Entire family units must be culled because of strong family bonds. 3,13,15 KNP culled elephants from 1967-1994, 12,29,30 but stopped after receiving pressure from international and animal welfare groups. 12 Bison can be shot in the field or sent to an abattoir. 20 Selection for bison culling is determined by desired percentages of age groups. 31 EINP has used culling since 1929 to control the bison population. 32 Meat from both elephant and bison culls have been sold or donated to surrounding communities. 13,20 Discussion and Management Implications KNP and EINP originally used a similar strategic approach of focussing on population size when applying population control methods, but KNP has now proposed to use  28 Thus, to manage impacts rather than specific population numbers it has been suggested to use fire and water management to guide elephants into the proposed zoned areas for high and low impacts. However, in KNP, cow/calf herd distribution is related to rivers more than artificial watering holes, whereas bulls appear to prefer artificial watering holes. 33 These distribution trends make it more complicated to control distribution. There are also the effects on other species to consider. The impacts of controlled burns on elephant habitat use are still in their infancy. It can be inferred from vegetation preferences that a freshly burned section would deter elephants because they prefer vegetation that is diverse and complex. 28 However, this strategy requires an understanding of the impacts of abiotic and biotic factors on the distribution of large herbivores 34 and how they can be used to mimic natural distribution patterns.
EINP has used the methods of culling and translocation, with variation in implementation. Currently, EINP uses a 'fast-tracking' method of translocation where only those bison leaving the park are processed in corrals. 20 Historically, with fairly high accuracy of bison numbers, almost all bison in the park were processed every 1-2 years. 32 EINP can benefit from integrating KNP's proposed impact management into their bison management strategy which could result in a better mimicry of natural processes. There is no water management plan in EINP like there is in KNP because of the different ecosystem. However, EINP has a fire and vegetation management plan that could be incorporated into the bison management plan. Bison have been found to prefer newly burned areas because of the high quality food, even in wooded areas. 34 Bison will naturally move into more freshly burned areas thereby decreasing their impacts on other areas. The integration of fire management into the bison management plan would assist EINP in forming a management strategy based on impact control rather than specific population numbers. Since EINP's management plan currently consists of what KNP has done in the past, there are no fundamental management ideas from EINP that can be integrated into KNP's current elephant management strategies. What KNP needs now is an approved management plan for their elephant population so that they can continue to support their rich biodiversity.
The choice of strategies to manage populations of large herbivores is deeply impacted by public opinions (i.e. social views on culling of elephants have delayed approval of KNP's elephant management plan, even though culling occurs for buffalo and other ungulates). To increase public acceptance, parks staff must provide the public with a rationale for using any strategies for managing large herbivore populations. In addition, animal populations are subject to fluctuations and chance events that can drastically alter population sizes. KNP's proposed elephant management plan appears to be trying to accommodate for these natural changes in elephant populations and use natural distribution factors, while still maintaining biodiversity.
With parks around the world increasingly becoming 'islands' of wilderness, management plans should start addressing impacts relating to population sizes of species since non-management approaches will no longer be effective. A comparison of large herbivore management strategies at KNP and EINP demonstrates how distant parks can learn from each other about ways of managing similar wildlife issues using the same methods but with different strategies guiding the implementation. As well it serves as a case study highlighting the impacts of large herbivores with abundant populations that is relevant to herbivore species of any body size. Such comparisons, resulting from a careful examination of underlying principles and practices, can be beneficial for other parks around the world in developing innovative wildlife management strategies, sometimes using methods they already employ.