Red List of South African Species

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Vulnerable (VU)
Assessors: Frans Radloff Coral Birss Carly Cowell Dean Peinke Desire Dalton Antoinette Kotze
Contributors: Jeanetta Selier
Reviewers: Graham Kerley Matthew Child David Mallon

Rationale (Changed due to New Information)

The Bontebok is a subspecies endemic to the East Coast Renosterveld bioregion within the Cape Floristic Region (CFR) of the Western Cape. It is an important South African conservation success story where a subspecies has been brought back from the brink of extinction in the 1830s by farmers in the Bredasdorp area. Currently (2014), the population estimate within the natural distribution range is 515 mature individuals (using a 75% mature population structure) on formally protected areas (including the Denel Overberg Test Range managed as an ecological unit within De Hoop Nature Reserve). Due to a lack of natural habitat availability within the natural range, the provincial conservation management authority, CapeNature, allows benign introductions of subpopulations outside of the historical range, in areas geographically adjacent to the natural range and possessing broadly similar habitat. Reintroductions into two such formally protected areas have been in place for at least five years with successful reproduction and these have increased the mature population to 664 individuals. Bontebok also exist on ranch lands and other private properties within the natural and benign introduction range. However, there are known incidences of intensive management and/or hybridisation with Blesbok (D. pygargus phillipsi) within these subpopulations. Preliminary estimates of privately owned subpopulations nationally, suggest that only 33–39% of these subpopulations can be considered sufficiently wild to be included in this assessment. This adds between 118 and 1,272 individuals (by extrapolating this proportion to the private properties with the most or least abundant subpopulations respectively), which potentially increases the mature population size to between 752 and 1,618 individuals. However, the overall numbers of pure Bontebok within this population range and the intensity of management of these subpopulations are currently unknown.

Although Bontebok numbers within the benign introduction and extra-limital ranges are increasing, the core population within the natural range has not increased since 2004 (770 individuals in formally protected areas in 2004 compared to 686 individuals in 2014). Protected area expansion possibilities are limited within the natural range, thereby limiting core population growth. Currently, the extent of occurrence within the natural range is estimated at 8,779 km² and the current observed area of occupancy is 602 km². Including all known Bontebok-containing areas within both the natural and benign introduction ranges yields 1,453 km² of observed occupancy. We infer a continuing loss of suitable habitat from ongoing agricultural and urban expansion within the CFR (within the Western Cape, 107 km² land was converted to agriculture per year between 2006 and 2011, 31% occurred within Critical Biodiversity Areas; and there has been an 8.6% increase, from 1,029 km² to 1,118 km², in urban expansion between 2000 and 2013).

Given that the estimated mature population size (within the natural and benign introduction areas) ranges from 514 to 1,618 individuals, that numbers of hybrid animals are unknown, and that no long-term data from the private sector exist to prove that the population has passed the threshold for Near Threatened for five years, we take a precautionary purview and list Bontebok as Vulnerable D1. Additionally, because the upper estimate of the potential area of occupancy (AOO) is 1,453 km² and that there is continuing loss of suitable habitat that limits population expansion within both the natural and benign introduction range, as well as that all subpopulations are fragmented by fencing with no metapopulation plan in place, Bontebok also qualifies for Vulnerable under criterion B.

The major threats to Bontebok are the uncertainty around the number of hybrids within the existing population, lack of habitat availability within its natural range (thus limiting population expansion), and the lack of a metapopulation plan to sustain genetic diversity. High incidences of hybridisation might render the majority of Bontebok subpopulations unfit for Red List inclusion and low genetic diversity may ultimately reduce the resilience of the subspecies. These threats should be counteracted through (1) the identification of all pure Bontebok subpopulations (and thus more accurate assessment of true population size), (2) the development and implementation of an active metapopulation management plan for the pure Bontebok subpopulations to simulate gene flow, prevent inbreeding and to sustain a flourishing and resilient population; and (2) exclusion of Blesbok from core areas supporting Bontebok inside its natural range. Conservationists should thus incentivise landowners to become Bontebok stewards to expand the conservation estate for this subspecies within the natural and benign introduction range. Bontebok are able to use a variety of habitats, the key features being grassy landscapes and the availability of water. For example, they readily utilise short grass areas and transformed landscapes. This tolerance and hence relative ease of management, together with their iconic status, facilitates stewardship prospects. This assessment should be revised when such data become available.

Distribution

Bontebok are endemic to the Western Cape, South Africa, although introductions have been made in most provinces. Historically, they were confined to the coastal plain (60–200 m) east of the Kogelberg in the Western Cape where they are believed to have concentrated on the renosterveld areas (Boshoff and Kerley 2001, Skead 2011). Early settlers may have confused Bontebok and Blesbok (D. pygargus phillipsi), which overestimated the historical distribution of Bontebok. In reality the subspecies were separated by at least 320 km at the time of European settlement (Skinner and Chimimba 2005, Boshoff et al. 2015). Here, overhunting reduced it from locally abundant to the verge of extinction. It was saved from extinction in the mid-19th century by a few Cape farming families who protected the small remnant subpopulations. From a low of 22 animals in the original Bontebok National Park (established near Bredasdorp in 1931), the Bontebok population has gradually recovered (Van Rensburg 1975). Translocated Bontebok from Bontebok National Park have formed the nucleus of reintroduced populations in other protected areas such as provincial and local authority nature reserves. The Bontebok National Park subpopulation is genetically pure (tested in April 2014[HDM1] ).

Suitable natural habitat within the indigenous natural range (IDR) is limited to the remaining renosterveld patches and some grassy micro-habitat patches in the fynbos areas of the Overberg region. The extent of occurrence is estimated to be 8,779 km², of which 3,664 km² comprises of remaining natural habitat (GeoTerraImage 2015), which can be construed as the maximum potential area of occupancy (AOO). However, only 623 km² of the preferred renosterveld habitat remains within the natural range, which closely matches the current estimated AOO within the IDR of 602 km². The latter may be an overestimate because only approximately 12% of De Hoop Nature Reserve is actually used by Bontebok (Radloff 2008), and Agulhas National Park currently has very little renosterveld within it. Thus, the amount of utilisable or optimal habitat is both limited and fragmented within the natural range. The AOO estimate increases to 1,153–1,453 km² depending on whether only protected areas, or also private properties are included within the extended natural range.

Preliminary genetic analyses indicate a low genetic variation within the IDR population (van der Walt et al. 2001, 2013). Within the IDR, the subpopulation is fragmented into small subpopulations restricted by fences. Habitat loss within the IDR has been so extensive that the area is estimated to be able to, at best, support 38% of the pre-transformation population potential (Kerley et al. 2003). The poor quality and limited availability of remaining habitat within the IDR necessitated the regulatory extension of their range, which we label here the benign introduction range (BIR) based on the latest Red List guidelines (IUCN Standards and Petitions Subcommittee 2014). This range extension has been spatially modelled to include habitat aspects such as altitude, slope and major preferred vegetation types, within a range which had been demarcated for regulatory purposes, whereby translocations of the subspecies were permitted outside its indigenous range prior to 2012. The BIR has enabled additional utilisation of this subspecies by private land owners and the creation of a buffer population from which to augment subpopulations within the IDR. Both West Coast and Table Mountain National Parks are located within the BIR. To date, however, no reintroductions into IDR protected populations have occurred. For example, Agulhas National Park only has two Bontebok but the long term plan is to source animals for future translocations from Table Mountain and West Coast National Parks. All translocations in the Western Cape are subject to the CapeNature Bontebok Conservation, Translocation and Utilisation Policy (Birss et al. 2013), which requires all proposed Bontebok for translocation to be genetically tested and declared pure before release. Some Bontebok subpopulations in the BIR can be considered ex situ conservation or benign introduction as they are within an environment that is reasonably close to the indigenous range and variables such as winter rainfall, disease, and habitat are considered similar enough to create comparable selection pressures. Additionally, this is a region where spatial separation from Blesbok is feasible, which has specific value in controlling the issuing of permits for Blesbok in this area. Importantly, the population within the IDR is <1,000 mature individuals and thus the BIR concept is a mechanism to support a viable population. However, at present, there is no metapopulation plan to connect subpopulations within the BIR to the IDR and thus the two populations are effectively isolated. Similarly, until genetic testing has been completed for both formally protected and privately protected subpopulations, it is uncertain which subpopulations should form part of the metapopulation. Thus, a precautionary approached is employed in including subpopulations.

Extra-limital subpopulations have been established on private farms or ranches in at least the Eastern Cape, Northern Cape, Free State and North West provinces, and there is pressure to increase the extent of introduction (Power 2014). These extra-limital introductions could be detrimental to the subspecies because Bontebok are adapted to very unique habitat and climatic conditions: the East Coast Renosterveld bioregion receives some rain throughout the year but has a distinct peak during winter with about 65% of rain falling between April and October. These animals thus adapted over at least 20,000 years in isolation from the Highveld-bound Blesbok to a climate of wet and cold winters and warm and dry summers. This is opposite to the conditions experienced by Blesbok that experience warm and wet summers and very cold but dry winters.

Population trend

Trend

Due to its restricted range, habitat transformation for agriculture, competition with domestic grazers and over-hunting, the Bontebok population reached a critical low in 1931 when 22 animals were fenced into the newly proclaimed Bontebok National Park (Van Rensburg 1975). Bontebok National Park was proclaimed in the Bredasdorp district with the specific goal of protecting a subpopulation of the subspecies (Barnard and van der Walt 1961). Population numbers increased, but remained below 100 individuals. It was then discovered that the animals at this site suffered from copper deficiency and high parasite infestation, specifically lungworm (Protostrogylus spp.). The park was also very small (6.8 km²) and was extensively covered (80%) by the unpalatable renosterbos (Dicerothamnus rhinocerotis) (Barnard and van der Walt 1961). In 1960, the present site of Bontebok National Park was proclaimed close to Swellendam and 84 Bontebok were translocated there, of which 61 survived (Penzhorn 1971). In the new area, numbers increased to 320 in 1981 with the current subpopulation maintained at around 250 and is currently 190 individuals after an annual off-take in 2014. The combined subpopulation for De Hoop Nature Reserve and neighbouring Denel Overberg Test Range, which is managed as an ecological unit, is currently estimated at 492 individuals, with a maximum of 526 individuals ever recorded. However, this subpopulation is prone to crashes if not managed correctly (Scott 1993). The current subpopulation size for Aghulhas National Park and Salmondsdam Nature Reserve is only two individuals, emphasising the urgent need for a metapopulation plan to create viable subpopulations. 
Based on field surveys and censuses, we can determine the current number of Bontebok with reasonable accuracy. Within the natural distribution range, there are 686 individuals in four formally protected areas (we include the greater De Hoop Area that includes De Hoop Nature Reserve and the privately owned Denel Overberg Test Range here). Subpopulation demographics from Bontebok National Park suggest a mature population structure of 75%, where a 1974 survey tallied 360 animals of which 71 were under two years of age and not deemed sexually mature (De Graaff et al. 1976). Using the figure of 75% thus implies that there are 515 mature individuals within the natural range. If the subpopulations existing on formally protected areas within the benign introduction range are considered, the total mature population size may be considered to be 664. Additionally, there are an observed 219 individuals on 50 private properties within the natural range, which increases the mature population size to 828 individuals. However, some of these subpopulations are intensively managed, or contain hybrids (Birss et al. 2013), and are thus not eligible for inclusion (IUCN Standards and Petitions Subcommittee 2014). For the latter, while both Bontebok National Park and Greater De Hoop Areas subpopulations have been genetically certified as pure, most private properties have not been tested and, of those tested, hybrids have been found (CapeNature unpubl. data). Ongoing testing will determine the number of subpopulations on private properties that are pure and thus eligible for inclusion in the Red List. For the former problem of intensive management, a preliminary analysis to determine which private subpopulations can be considered wild, reveal that 33–39% of private subpopulations are eligible for inclusion in the assessment (based on a sample of N = 18 private subpopulations from across the country, Endangered Wildlife Trust unpubl. data). Depending on whether we extrapolate this proportion to the private farms within the natural and benign introduction range (N = 141 properties) with the most or least abundant subpopulations yields an additional 118–1,272 individuals respectively, and thus 89–954 mature individuals. This increases the mature population range, including all formally protected areas and private subpopulations within both the natural and benign introduction range, to 752–1,618 individuals. However, it is important to note that this estimate does not include the serious threat of hybridisation which might reduce the number of eligible private subpopulations dramatically. The crux of Bontebok conservation is determining genetic purity of its subpopulations and the consensus among the assessors is that population size must be conservatively based on only genetically tested subpopulations that have been shown to be pure.

Overall, the core population within the natural range on protected areas has not increased since the 2004 estimate (Friedmann and Daly 2004), where the total population tallied to 770 individuals compared to 686 individuals in 2014. Outside of the natural and benign introduction range, however, Bontebok subpopulations are increasing. There are an observed 4,857 individuals (existing in extra-limital areas, which are not included in this assessment (IUCN Standards and Petitions Subcommittee 2014). This is probably an underestimate as not all information from the private sector is available. Extra-limital subpopulations are not considered to possess conservation value for reintroduction or supplementation in the natural range. For example, although the Tsolwana Nature Reserve, Eastern Cape, subpopulation was registered as pure Bontebok based on the phenotypic but not genetic assessment (Fabricius et al. 1989), it is not considered an ideal source for reintroductions because the selection pressure outside the natural range may differ significantly. Here, the number of years/generations since departure from the natural range will be important: if it is only a couple of years or generations since departure it might still be an acceptable source for reintroduction providing that the subpopulation was not subjected to selective breeding (Champagnon et al. 2012). Initial indications are that 116 extra-limital subpopulations are from herds that have been registered as pure based on phenotypic traits and another 229 are from herds that have been genetically tested (D. Dalton unpubl. data). These data still need more analysis though as there are a number of owners listed where numbers are not given and it is suspected that a greater proportion may also have been tested and or registered. However, these subpopulations, even if pure, are still of limited conservation value due to the problems associated with re-stocking from extra-limital subpopulations and out-breeding depression (Champagnon et al. 2012). Thus, extra-limital subpopulations of Bontebok, especially if maintained over several generations are of little to no value to Bontebok conservation.

A metapopulation management plan is desperately needed to sustain Bontebok genetic purity and diversity. Within the natural range, there are many small properties containing small subpopulations which cannot increase in size because they are limited by space. In the natural areas, there is a space limitation due to extensive crop production. For example, the average subpopulation size on private properties in the Western Cape is 14 ± 20 individuals (N = 127 properties), compared to 38 ± 36 individuals in the Eastern Cape (N = 69 properties), 28 ± 31 individuals in the Free State (N = 22 properties), and 46 ± 49 individuals in the Northern Cape (N = 27 properties). Of 6,677 individuals estimated to have potentially occurred in the natural range before habitat transformation, only 2,544 individuals are estimated to be able to be supported now (Kerley et al. 2003). Thus, although there is still scope for the core population to increase (currently only 905 individuals in total within the natural range), protected areas and private conservation areas must significantly expand to create larger areas of natural habitat for the population and these should be connected to a wider metapopulation that includes the benign introduction range to become a resilient population.

Threats

The major threats for this subspecies are hybridisation with Blesbok, the lack of available habitat within its natural range (thus limiting population expansion), low genetic diversity, which was probably the result of two bottlenecks caused by overhunting and disease (van der Walt et al. 2001) and poor gene flow between subpopulations. Habitat loss is severe and in some cases irreversible. Although the population size of what we hope is pure Bontebok has increased, it has remained relatively low compared to other ruminants (for example, Broders et al. 1999). The ultimate threat is the indiscriminate and uncontrolled movement of Blesbok, Bontebok and Blesbok/Bontebok hybrids across the country. As such, the exact status of Bontebok and the long term security of the subspecies is still uncertain. Additionally, there is uncertainty over the impact climate change will have, especially in terms of the synergistic threat of habitat loss. The corresponding threats are: 
  1. Habitat loss and consequent lack of available habitat, leading to fragmented and isolated subpopulations, which exist primarily on suboptimal habitat. Here Bontebok are currently managed as sources for properties outside the natural range for commercial gain, while comparatively little effort is being directed towards habitat rehabilitation and restoration to secure the future of the core Bontebok population within the natural range in the Western Cape. Currently, only 9–12% of renosterveld remains (Von Hase et al. 2003, Cowell and Birss 2013). The opportunity exists for fragmented natural areas to be linked via corridors to larger natural areas and to conserve more of the threatened renosterveld vegetation.
  2. Low genetic diversity within the core natural range population and thus lack of resilience to environmental change. The historical bottlenecks are currently exacerbated by the fragmented and isolated nature of the subpopulations, which can lead to decreased genetic variation within individual subpopulations due to genetic drift. Additionally, biased selection pressures (for coat colour, pattern and horn sizes) further threaten the genetic integrity and viability of the population with inbreeding depression. The decision to translocate pure Bontebok, as part of a metapopulation should therefore not only consider the genetic purity of the subspecies, but also consider the genetic relationships among the subpopulations involved.
  3. Hybridisation with Blesbok, which has been introduced into the Western Cape, may affect a significant number of subpopulations. Hybridisation between the two has occurred due to human-mediated mixing of the separate subspecies. However, what is not known is the full extent to which hybridisation occurs. Preliminary research shows that a high proportion of Bontebok could be hybrids (van Wyk et al. 2013). Hybridisation was detected in 33% (40 of 121) of the samples with unknown purity. It is thus vital that hybridisation tests be expanded to include all regions in the natural and benign introduction ranges where potentially pure Bontebok subpopulations may be under threat. Negative consequences associated with hybridisation include reduction of fitness, alteration in the genetic structure of populations and the interference of locally co-adapted gene complexes (Allendorf et al. 2001). However, a study assessing body condition of hybrids in Oviston Nature Reserve, Eastern Cape, found that hybrid body condition was similar to Blesbok within their natural range and is probably adequately adapted to the Eastern Cape environment (Schmidt 1999). Further research is needed for subpopulations in the Western Cape. In the worst case scenario hybridisation might lead to the extinction of both the Blesbok and Bontebok subspecies, before the debate on the taxonomic status of the two subspecies/species has been resolved.
  4. The overall management concern is the lack of a metapopulation management strategy for this subspecies, without which there is mismanagement and misunderstanding between stakeholders. The development of a strategic management plan that is aimed at ensuring the long term survival of Bontebok and uniting stakeholders is urgently required. Such a strategy should include incentives for landowners to continue managing and conserving Bontebok and it should build on the strengths of the previous success of saving this subspecies when only approximately 22 individuals remained in the 1930s.

Uses and trade

The horns and skins are traded as part of trophy hunts, and live animals are traded at game auctions. Individuals are utilised from captive breeding, ranched (extensive) and wild (formally protected) areas. Activities on private land are mainly for ecotourism, trophy hunting and breeding of trophy animals. As such, the subspecies is widely hunted and traded outside of its natural range where it has increased dramatically in numbers but unfortunately have limited to no conservation value. The translocation of Bontebok and Blesbok outside their natural range is a significant threat to the continued existence of the two subspecies, as hybridisation takes place and becomes increasingly likely. Additionally, there is a concern that there is a net flow of individuals away from the natural range. Western Cape permit data for the last 10 years show that almost 600 animals have been translocated from the natural range and a further 200 from the benign introduction range. Since hunting is not regulated through direct permitting, the number of hunted Bontebok is unknown. Bontebok is a subspecies listed on Certificates of Adequate Enclosures (CoAEs), which means that harvest/hunt/export is not regulated but exempted from separate applications and consequently there is limited regulatory control of subpopulations on private properties (Birss et al. 2013).

At present there has been no reported illegal offtake of Bontebok in any of the national parks or provincial parks where they occur. Given the limited genetic diversity of Bontebok, a strict control of live animal sales needs to be set in place to limit inbreeding in the long term. Additionally, exporting animals outside of the natural range can deplete the genetic pool of the core population and management plans should thus ensure that the translocation of animals, as part of a managed metapopulation, is kept between the natural and benign introduction range.

Farmers saved this subspecies from extinction but modern wildlife ranchers have ambiguous effects on Bontebok conservation. Wildlife ranching is not prominent in the natural range and it appears as if the largely extra-limital industry is impacting negatively on landowners within the natural range who want to contribute to the conservation of the subspecies but increasingly cannot afford to buy and stock Bontebok. The value of the subspecies has increased in the last five years and there is an increasing demand for extra-limital introductions such as in the North West Province (Power 2014). Genetic testing for purity has increased the price and the demand for pure animals. This might act as an incentive to not only conserve the subspecies itself, but also its preferred habitat, the renosterveld, although there is no evidence that this is currently happening in its native range. Prudent veld management is needed in the native range where Bontebok do occur, as improving grass cover within renosterveld might result in the loss of plant species diversity, including species classified as rare or threatened (Novellie and Kraaij 2010). On the negative side, there is evidence that some private landowners deliberately hybridise Bontebok and Blesbok for trophy hunting purposes (Schmidt 1999), and artificial selection for colour morphs and exceptional horn lengths is suspected. The hybrid and artificial selection threat must be curbed through a management plan taken up and enforced by all participating landowners.

Conservation

Currently, key protected areas include Bontebok National Park (genetically certified pure subpopulation within the natural range); Table Mountain National Park (genetically certified pure subpopulation within the benign introduction range); De Hoop Nature Reserve and Denel Overberg Test Range (largest subpopulation within the natural range); and Agulhas National Park (potential for significant subpopulation growth within the natural range). Although subpopulations have exhibited positive growth rates, offtakes and translocations from within the natural range have suppressed overall population growth. Similarly, the lack of available habitat within the natural range has inhibited population expansion and an increase in area of occupancy. The net effect is that the core Bontebok population has not increased within the natural range since the 2004 assessment (Friedmann and Daly 2004). To redress this, the main interventions should be the development and implementation of a Biodiversity Management Plan and metapopulation strategy, as well as the acquisition and connection of additional natural habitat within the natural and benign introduction distribution range.
  1. Biodiversity Management Plan: Regulation of translocation is required to prevent hybridisation with Blesbok and Bontebok/Blesbok hybrids. Subpopulations within the natural and benign introduction ranges should be tested for purity and flagged as important subpopulations to potentially reintroduce or augment other areas. During a National Workshop on Bontebok Purity Testing and Management held in 2009, the National Zoological Gardens of South Africa was mandated to further develop a scientifically defensible genetic test for Bontebok and Blesbok hybridisation using microsatellite (DNA) markers. The genetic test developed is supported by peer-reviewed scientific publications and forms the basis for the updated CapeNature Bontebok Conservation, Translocation and Utilization Policy, as well as the national implementation of a standardised genetic testing protocol (Birss et al. 2013). Any animal which will be translocated must be tested and must be fitted with a microchip, and all test results are stored in a central database at the National Zoological Gardens. This information will inform metapopulation management, but will also identify properties where hybrids have been identified, which will allow conservation agencies to enforce regulations. It is foreseeable that collaboration between provinces will identify which subpopulations need to be targeted for further testing. Thus far, legislation appears to be working to prevent hybridisation. The CapeNature policy instructs that hybrids be destroyed within 48 hours of results being known and thus far, destruction orders have been carried out effectively. Once subpopulations have been identified that are eligible for inclusion in the Red List, a metapopulation plan should be employed to ensure translocations are used to sustain genetic diversity and increase the number and size of subpopulations within the natural and benign introduction ranges. The superior financial value of pure Bontebok subpopulations may be one mechanism to incentivise landowner participation.
  2. Securing natural habitat: Conservancies and corridors must be created to conserve areas of suitable remaining natural habitat to ensure positive population growth for Bontebok in the natural range. For this to happen, the Biodiversity Management Plan currently being drafted must be adopted by public and private stakeholders, and biodiversity stewardship schemes should be strategically established. The importance of biodiversity stewardship is clear: currently, Agulhas National Park only has two Bontebok. However, 38 Bontebok from the Greater De Hoop Area were donated to the Nuwejaars Special Management Area (NSMA) in 2010. The NSMA is situated on the southern Agulhas Plain which includes a variety of threatened vegetation types, amongst which are Overberg Sandstone, Overberg Sand Fynbos, Overberg Dune Strandveld, Agulhas Limestone Fynbos, Elim Ferricrete Fynbos, De Hoop Limestone Fynbos and Central Rûens Shale Renosterveld. The NSMA consists of landowners who are committed to contributing to and pioneering this landscape conservation initiative, while maintaining their farming operations, and highlights the stewardship potential of private landowners for Bontebok conservation. Expansion of existing protected areas until sufficient habitat for at least 2,000 Bontebok has been achieved within the natural range and creating links between protected areas wherever possible, is recommended (Kerley et al. 2003). Conservationists and private landowners should also restore renosterveld areas in the natural range and promote other mechanisms of slowing land-use change in the area. For example, offsets can be put in place where new land is being farmed.
Bontebok conservation is primarily concerned with growing the population to achieve viability (Traill et al. 2007) through increasing available habitat, maintaining metapopulation processes to offset inbreeding and preventing hybridisation. Overall, more effort should be invested in connecting, rehabilitating and sustaining a resilient and genetically pure population within the natural range, which can then be used to seed private properties wanting to maintain pure subpopulations. Conservationists should concentrate on rehabilitating renosterveld habitat within the natural range (Von Hase et al. 2003, Cowell and Birss 2013), and incentivising landowners to become Bontebok stewards to expand the conservation estate for this subspecies in natural and benign introduction range. This subspecies evolved in an environment very different from the Highveld grassland conditions associated with Blesbok and has the potential to act as a flagship subspecies for the renosterveld, thus incentivising the conservation of this highly threatened veld type. Additionally, Bontebok are tolerant to human activities and adapt to changes in the landscape. They readily utilise short grass areas and transformed landscapes, thus facilitating stewardship opportunities. We recommend that Bontebok outside the natural and benign introduction range should not be used to reintroduce or augment subpopulations within these ranges, and that conservation efforts should be focused on securing and growing the native subpopulations that have been experiencing the most natural selection pressures. Although Bontebok are genetically depauperate, genetic recovery through hybridisation is not recommended given the potential alpha level taxonomic distinction between the Blesbok and Bontebok (van der Walt et al. 2013). We recommend that the putative subspecies be managed as separate evolutionary significant units and that more research is undertaken to determine the taxonomic statuses of Bontebok and Blesbok.

Recommendations for land managers and practitioners:
  • At present there is no formal management plan for Bontebok in any province and SANParks currently has no specific management strategy. Bontebok National Park, which was established to ensure the survival of Bontebok, has the largest subpopulation within SANParks and can no longer expand in size. The Agulhas National Park has been identified as ideal habitat to establish a subpopulation of Bontebok but the park currently has no fencing. Funding has been requested to erect fences and only then can Bontebok be augmented here. Conservation agencies must identify similar sites for subpopulation augmentation and incorporate the sites into a metapopulation plan. SANParks and CapeNature have initiated a process to develop a Biodiversity Management Plan (BMP) for Bontebok. The BMP will be aimed at ensuring its long term survival in the natural and benign introduction range. The management plan should prioritise the identification of pure Bontebok subpopulations and make recommendations for the alignment of existing legislation to achieve the desired outcomes of the BMP. National and provincial legislation must be aimed at policies that prevent the co-occurrence of Blesbok and Bontebok and the exclusion of Blesbok within the natural distribution range of Bontebok. It should also provide guidelines concerning the keeping of Blesbok and Bontebok in close proximity to each other in extra-limital areas. Guidelines on the movement and mixing of subpopulations inside and outside the natural, benign introduction and extra-limital ranges should also be created and implemented.
  • Genetic testing, using the microsatellite markers identified by van Wyk et al. (2013), must be completed for all putative metapopulation subpopulations within the natural and benign introduction ranges.
  • Harvesting to prevent overstocking/grazing can help maintain the remaining renosterveld habitat. There is an opportunity to use Bontebok as a flagship subspecies for the conservation and rehabilitation of renosterveld. However, if Bontebok become common outside its natural distribution range, the financial incentive associated with its rarity will be lost and with it the opportunity to use the subspecies to promote renosterveld conservation. This emphasises the importance of implementing the BMP mentioned above. 
  • Extra-limital subpopulations are expected to have little conservation value due to the increased probability of hybridisation with Blesbok, as well as the very different environmental conditions, and consequent selection pressure, they are exposed to. Restocking or reintroducing individuals from such subpopulations into natural or benign introduction subpopulations could cause deleterious effects such as changes in behaviour, morphology and demography that could in turn erode local adaptation and decrease the fitness of the core population. Thus, reintroduction or supplementation of individuals from extra-limital subpopulations is discouraged.
Research priorities:
  • It is of utmost importance that the taxonomic status of Bontebok and Blesbok, as either subspecies of each other or separate species, is resolved as a matter of urgency. The uncertainty around their status thwarts decision-making and consequent conservation efforts. 
  • The size, demography, location and health status of pure Bontebok subpopulations need to be identified so that a metapopulation management plan can be designed and implemented.
  • A recent survey for farms which had Bontebok in the Western Cape in the past revealed that many properties no longer have Bontebok and anecdotal information suggests that animals die from contracting pathogens from domestic livestock (lungworms) and from poor habitat quality. Understanding which private properties have pure Bontebok with stable or increasing populations and could be incorporated as core sites into the metapopulation is crucial.
Encouraged citizen actions:
  • Landowners should form conservancies within the natural and benign introduction ranges to support free-roaming Bontebok herds and keep their herds genetically pure or cooperate with provincial conservation authorities to translocate according to a metapopulation plan. 

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 title = {Historical incidence of the larger land mammals in the broader Western and Northern Cape provinces},
 year = {2011}
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,@techreport{3326,
 address = {Pretoria},
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 year = {2015}
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,@techreport{5228,
 author = {Power, R.J.},
 institution = {Department of Economic Development, Environment, Conservation & Tourism, North West Provincial Government, Mahikeng},
 title = {The distribution and status of mammals in the North West Province},
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,@book{6236,
 author = {IUCN Standards and Petitions Subcommittee},
 title = {Guidelines for Using the IUCN Red List Categories and Criteria. Version 11},
 year = {2014}
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,@article{6236,
 author = {Beukes, P.C.},
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 pages = {103--108},
 title = {Responses of grey rhebuck and bontebok to controlled fires in coastal renosterveld},
 volume = {17},
 year = {1987}
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,@book{6236,
 author = {Friedmann, Y. and Daly, B.},
 publisher = {CBSG Southern Africa, IUCN SSC Conservation Breeding Specialist Group, Endangered Wildlife Trust, South Africa},
 title = {Red Data Book of the Mammals of South Africa: A Conservation Assessment.},
 year = {2004}
}
,@phdthesis{6236,
 author = {Radloff FGT},
 school = {University of Stellenbosch, Stellenbosch, South Africa},
 title = { The ecology of the large herbivores native to the coastal lowlands of the Western Cape, South Africa},
 year = {2008}
}
,@article{6236,
 author = {Allendorf, F.W., Leary, R.F., Spruell, P. and Wenburg, J.K.},
 journal = {Trends in Ecology & Evolution},
 pages = {613--622},
 title = {The problems with hybrids: setting conservation guidelines},
 volume = {16},
 year = {2001}
}
,@article{6236,
 author = {Barnard PJ, van der Walt DK},
 journal = {Koedoe},
 pages = {105--109},
 title = {Translocation of the bontebok (<i>Damaliscus pygargus</i>) from Bredasdorp to Swellendam},
 volume = {4},
 year = {1961}
}
,@phdthesis{6236,
 author = {Beukes, P.C.},
 school = {University of Stellenbosch},
 title = {Sommige aspekte van die ekologie van die Vaalribbok (<i>Pelea Capreolus</i>, Forster 1790) in die Bontebok Nasionale Park},
 year = {1984}
}
,@article{6236,
 author = {Bigalke, R},
 journal = {Fauna Flora},
 pages = {95--116},
 title = {The bontebok (<i>Damaliscus pygargus</i> (Pall.)) with special reference to its history and preservation},
 volume = {6},
 year = {1955}
}
,@techreport{6236,
 address = {Cape Town},
 author = {Birss C, van Deventer JD, Hignett DL, Brown C, Gildenhuys P, Kleinhas D},
 institution = {Western Cape Nature Conservation Board, CapeNature},
 title = {Bontebok conservation, translocation and utilisation policy. Version 1},
 year = {2013}
}
,@article{6236,
 author = {Boshoff, A.F. and Kerley, G.I.H.},
 journal = {African Zoology},
 pages = {245--273},
 title = {Potential distributions of the medium-to large-sized mammals in the Cape Floristic Region, based on historical accounts and habitat requirements},
 volume = {36},
 year = {2001}
}
,@article{6236,
 author = {Boshoff AF, Landman M, Kerley G},
 journal = {Transactions Royal Society of South Africa},
 pages = {1--65},
 title = {Filling the gaps on the maps: historical distribution patterns of some larger mammals in part of southern Africa},
 volume = {70},
 year = {2015}
}
,@article{6236,
 author = {Broders HG, Mahoney SP, Montevecchi WA, Davidson WS},
 journal = {Molecular Ecology},
 pages = {1309--1315},
 title = {Population genetic structure and the effect of founder events on the genetic variability of moose, <i>Alces alces</i>, in Canada},
 volume = {8},
 year = {1999}
}
,@article{6236,
 author = {Champagnon J, Elmberg J, Guillemain M, Gauthier-Clerc M, Lebreton J-D},
 journal = {Journal of Nature Conservation},
 pages = {231--241},
 title = {Conspecifics can be aliens too: A review of effects of restocking practices in vertebrates},
 volume = {20},
 year = {2012}
}
,@techreport{6236,
 address = {Cape Town, South Africa},
 author = {Cowell C, Birss C},
 institution = {Cape Research Centre, South African National Parks},
 title = {Resource Information for Bontebok (<i>Damaliscus pygargus pygargus</i>) in South Africa. CRC-11-2013},
 year = {2013}
}
,@article{6236,
 author = {David JHM},
 journal = {Zeitschrift für Tierpsychologie},
 pages = {38--107},
 title = {The behaviour of the Bontebok, <i>Damaliscus dorcas dorcas</i>, (Pallas 1766), with special reference to territorial behaviour},
 volume = {33},
 year = {1973}
}
,@inbook{6236,
 address = {Amsterdam, The Netherlands},
 author = {},
 booktitle = {<i>Damaliscus pygargus</i>},
 editor = {David, J. and Lloyd, P.},
 publisher = {Academic Press},
 title = {The Mammals of Africa},
 year = {2013}
}
,@article{6236,
 author = {De Graaff G, van der Walt PT, van Zyl LJ},
 journal = {Koedoe},
 pages = {67--74},
 title = {Populasie-samestelling van die bontebok <i>Damaliscus dorcas dorcas </i>in die Bontebok Nasionale Park gedurende Januarie 1974},
 volume = {19},
 year = {1976}
}
,@article{6236,
 author = {Fabricius C, van Hensbergen HJ, Zucchini W},
 journal = {South African Journal of Wildlife Research},
 pages = {61--66},
 title = {A discriminant function for identifying hybrid bontebok x blesbok populations},
 volume = {19},
 year = {1989}
}
,@article{6236,
 author = {Kerley GI, Pressey RL, Cowling RM, Boshoff AF, Sims-Castley R},
 journal = {Biological Conservation},
 pages = {169--190},
 title = {Options for the conservation of large and medium-sized mammals in the Cape Floristic Region hotspot, South Africa},
 volume = {112},
 year = {2003}
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,@article{6236,
 author = {Kraaij, T. and Novellie, P.A.},
 journal = {African Journal of Range & Forage Science},
 pages = {21--27},
 title = {Habitat selection by large herbivores in relation to fire at the Bontebok National Park (1974–2009): the effects of management changes},
 volume = {27},
 year = {2010}
}
,@phdthesis{6236,
 author = {Luyt ED},
 school = {University of Stellenbosch},
 title = {Models of Bontebok (<i>Damaliscus pygargus pygargus</i>, Pallas 1766) habitat preferences in the Bontebok National Park and sustainable stocking rates},
 year = {2005}
}
,@article{6236,
 author = {Midgley GF, Hannah L, Millar D, Rutherford MC, Powrie LW},
 journal = {Global Ecology and Biogeography},
 pages = {445--451},
 title = {Assessing the vulnerability of species richness to anthropogenic climate change in a biodiversity hotspot},
 volume = {11},
 year = {2002}
}
,@article{6236,
 author = {Novellie, P.},
 journal = {Koedoe},
 pages = {1--17},
 title = {Interrelationships between fire, grazing and grass cover at the Bontebok National Park},
 volume = {30},
 year = {1987}
}
,@article{6236,
 author = {Novellie P, Kraaij T},
 journal = {Koedoe},
 pages = {1--5},
 title = {Evaluation of <i>Themeda triandra</i> as an indicator for monitoring the effects of grazing and fire in the Bontebok National Park},
 volume = {52},
 year = {2010}
}
,@article{6236,
 author = {Penzhorn BL},
 journal = {Koedoe},
 pages = {145--159},
 title = {A summary of the re-introduction of ungulates into South African National Parks (to 31 December 1970)},
 volume = {14},
 year = {1971}
}
,@article{6236,
 author = {Schmidt AG},
 journal = {South African Journal of Wildlife Research},
 pages = {145--147},
 title = {Body condition in blesbok x bontebok hybrids in the Oviston Nature Reserve, Eastern Cape Province},
 volume = {29},
 year = {1999}
}
,@phdthesis{6236,
 author = {Scott HA},
 school = {Port Elizabeth Technikon},
 title = {An investigation into possible reasons for a decline in numbers of bontebok <i>Damaliscus dorcas dorcas</i> (Pallas, 1766) in the De Hoop Nature Reserve, Southwestern Cape},
 year = {1993}
}
,@article{6236,
 author = {Traill LW, Bradshaw CJ, Brook BW},
 journal = {Biological conservation},
 pages = {159--166},
 title = {Minimum viable population size: a meta-analysis of 30 years of published estimates},
 volume = {139},
 year = {2007}
}
,@article{6236,
 author = {van der Walt, J.M., Nel, L.H. and Hoelzel, A.R.},
 journal = {Molecular Ecology},
 pages = {1679--1688},
 title = { Characterization of major histocompatibility complex DRB diversity in the endemic South African antelope <i>Damaliscus pygargus</i>: a comparison in two subspecies with different demographic histories},
 volume = {10},
 year = {2001}
}
,@article{6236,
 author = {van der Walt J, Nel LH, Hoelzel AR.},
 journal = {Conservation Genetics},
 pages = {243--248},
 title = {Differentiation at mitochondrial and nuclear loci between the blesbok (<i>Damaliscus pygargus phillipsi</i>) and bontebok (<i>D. p. pygargus</i>): implications for conservation strategy},
 volume = {14},
 year = {2013}
}
,@article{6236,
 author = {van Rensburg APJ},
 journal = {Koedoe},
 pages = {165--190},
 title = {Die geskiedenis van die Nasionale Bontebokpark, Swellendam},
 volume = {18},
 year = {1975}
}
,@article{6236,
 author = {van Wyk AM, Kotzé A, Randi E, Dalton DL},
 journal = {Conservation Genetics },
 pages = {589--599},
 title = {A hybrid dilemma: a molecular investigation of South African bontebok (<i>Damaliscus pygargus pygargus</i>) and blesbok (<i>Damaliscus pygargus phillipsi</i>)},
 volume = {14},
 year = {2013}
}
,@phdthesis{6236,
 author = {van Zyl LJ},
 school = {University of Stellenbosch},
 title = {Die waterbehoeftes en drinkgewoontes van die Bontebok <i>Damaliscus dorcas dorcas</i> (Pallas, 1766) in die Bontebok Nasionale Park},
 year = {1978}
}
,@techreport{6236,
 address = {Kirstenbosch, Cape Town},
 author = {von Hase A, Rouget M, Maze K, Helme N},
 institution = {Botanical Society of South Africa},
 title = {A finescale conservation plan for Cape lowlands renosterveld. Technical Report},
 year = {2003}
}
,@article{6236,
 author = {Watson LH, Kraaij T, Novellie P},
 journal = {South African Journal of Wildlife Research},
 pages = {158--166},
 title = {Management of rare ungulates in a small park: habitat use of Bontebok and Cape Mountain Zebra in Bontebok National Park assessed by counts of dung groups},
 volume = {41},
 year = {2011}
}
,@inbook{6236,
 author = {},
 booktitle = {Order Artiodactyla},
 editor = {Grubb P},
 publisher = {Smithsonian Institution Press},
 title = {Mammal Species of the World: A Taxonomic and Geographic Reference, 2nd edition.},
 year = {1993}
}
,]
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