Distribution
The Cape Mountain Zebra is endemic to the Cape Floristic Region of South Africa, and occurs in the Nama Karoo, Succulent Karoo, and Grassland Biomes (Skead 2007, 2011; Boshoff et al. 2015) (see Table 1 and Figure 1 in the Supporting Information). Its historical distribution extended throughout the great escarpment range in the Cape, south of the Orange River, including the Cape Fold Belt Mountains (the southern parts of the current eastern Western Cape Province), and the southern extent of the Northern Cape province (Figure 2 in the Supporting Information). Thus, although once widely distributed throughout the mountainous regions of the Cape, over-hunting and agricultural expansion had reduced the population to fewer than 80 individuals located in just five areas of the former Cape province by the 1950s (Millar 1970). Only three remnant subpopulations from the former natural distribution survived: Mountain Zebra National Park, Kammanassie and Gamkaberg Nature Reserves (Smith et al. 2008). It is postulated that in historical times they were separated from Hartmann’s Mountain Zebra (Equus zebra hartmannae; which occur mainly in Namibia), by an area devoid of mountainous habitat, the Knersvlakte, which separates the Kamiesberg in the north from the Roggeveldberge in the south (Novellie et al. 2002). However there are no historical (pre-1920) records of Hartmann’s Mountain Zebra south of the Orange River (Skead 2011).
The current distribution is limited to (at least) 75 fenced and isolated subpopulations spread throughout the former range. Subpopulations have been reintroduced to, amongst others, Karoo, Addo Elephant, Bontebok, Tankwa Karoo and Camdeboo national parks, De Hoop Nature Reserve, Commando Drift Nature Reserve, Baviaanskloof Wilderness Area and Tsolwana Nature Reserve (Hrabar and Kerley 2015). Two of these reserve populations (Commando Drift and Tsolwana) are possibly extralimital, as there are no historical records of the species east of the Great Fish River (Skead 2007). The subpopulation at Gariep Dam Nature Reserve in the Free State Province is significantly extra-limital (Boshoff and Kerley 2013) and not included in this assessment. Novellie et al. (2002) regarded the West Coast National Park as being within the historical range but there is some doubt about this given that it is on the coast and 70 km from the closest historically-recorded subpopulation in Picketburg (Skead 2011). The issue of whether to include West Coast National Park to be within the subspecies’ range has not been resolved (Figure 1 in the Supplementary Information). The subpopulation at Oorlogskloof Nature Reserve in the Northern Cape lies in a historical range of Cape Mountain Zebra. It was founded in 2003 with six males and 11 females from Gariep Dam Nature Reserve. In 2010, 18 animals (five males, 13 females) from Bontebok National Park, that appeared to be free of sarcoids, were translocated to Oorlogskloof Nature Reserve (Zimmermann et al. 2010). A number of Cape Mountain zebra have also been translocated to private properties located significantly outside of the natural range (for example, in the Free State and Northern Cape provinces) but these animals are not included in this assessment. There is no need to introduce this subspecies outside its natural range for conservation purposes.
In 2014/15, 66% of the area of occupancy (AOO) was constituted of formally protected areas (5,625 km²) and 34% private areas, which yielded a total AOO of 8,566 km² (Hrabar and Kerley 2015). Although formally protected areas have not increased in number, many have increased in size since 2009 (for example, Anysberg NR increased by 1,200 km² in 2012 and Tankwa Karoo NP has increased by 354 km²), thereby resulting in a 20% increase in formally protected habitat. Despite this expansion of protected areas and the rapid growth of the private sector contribution (including biodiversity stewardship sites), Cape Mountain Zebra habitat is likely to remain severely fragmented due to game fencing. Translocations between subpopulations as part of a metapopulation plan can potentially reduce the impacts of this fragmentation (such as loss of genetic diversity), but this does not take place consistently enough (Hrabar and Kerley 2015). Thus, further reintroductions, to both formally and privately protected areas, should be facilitated by a biodiversity and metapopulation management plan and follow the International Union for the Conservation of Nature (IUCN) guidelines on reintroductions (IUCN/SSC 2013).
Population trend
Trend
Cape Mountain Zebras were once widespread and numerous. Hunting and habitat loss to agriculture reduced them to just 80 individuals remaining in three relict populations in the 1950s (Bigalke 1952, Millar 1970): the Mountain Zebra National Park (MZNP) subpopulation consisted of 19 individuals, and the Kammanassie Nature Reserve and Gamka Nature Reserve subpopulations consisted of no more than five and six individuals at their respective nadirs (Millar 1970, Lloyd 1984). The total population therefore bottlenecked at around 30 individuals at this time. Subsequently, two thirds of the subspecies genotypic variation is located in just two of the subpopulations (Kamannassie and Gamkaberg Nature Reserves); all other subpopulations (except one, De Hoop Nature Reserve) originate from MZNP alone. Active metapopulation management is thus needed, and always will be needed, to ensure genetic diversity. Mountain Zebra National Park (MZNP) has provided founder individuals for around 30 subpopulations (Novellie et al. 2002), which has led to the undesirable state of over 91% of the genetic variability in the metapopulation deriving from one stock (Moodley and Harley 2005). Individuals have thus far not been removed from the other two remnant subpopulations, Kammanassie and Gamkaberg Nature Reserves, as they are currently too small, nor have these populations been supplemented with MZNP stock. Only De Hoop Nature Reserve, Western Cape has been founded with individuals from two of the original subpopulations (MZNP and Kammanassie), making it an important genetic source (Moodley and Harley 2005). However, the subpopulation at De Hoop Nature Reserve has experienced an annual decline of 6.6% (1995–1999) to 4.5% (1999–2005) (Smith et al. 2008). Overall, though, the combined population is increasing. From 1985 to 1995 the annual rate of increase was reported as 8.6 % (Novellie et al. 1996), 9.6 % between 1995 and 1998 (Novellie et al. 2002), 8.33 % between 2002 and 2009 (Hrabar and Kerley 2013) and 9.16% between 2009-2015 (Hrabar and Kerley 2015). However, estimating annual growth rate trends is challenging because most subpopulations are actively managed (maintained at sustainable stocking levels) and animals are often removed or augmented, which masks true growth rate.
Currently (2014/15), there are estimated to be between 1,714 and 3,247 mature individuals (using a 55% and 75% mature population structure respectively; Table 2 in the Supplementary Information). In formally protected areas alone, there are an observed 1,714–2,338 mature individuals (Table 2 in the Supplementary Information). Tankwa Karoo National Park, Western Cape is included as it is within the natural distribution range. Similarly, Oorlogskloof Nature Reserve in the Northern Cape Province is included as it within the historical range (Boshoff et al. 2015). A preliminary analysis to determine which private subpopulations can be considered wild, revealed that 81–98% of individuals on private land are eligible for inclusion in the assessment (N = 21 properties, A. Taylor unpub. data), which corresponds to “Extrapolated eligible total” in Table 2. This brings the total current population size within the natural distribution range to 1,582–2,157 mature individuals. Similarly, only including subpopulations with 50 individuals or more (“Total effective subpopulations”) yields 1,973–2,691 mature individuals (Table 2 in the Supplementary Information). Hybridisation with Plains Zebra (Equus quagga) has been identified as an emerging threat, where currently 28% of the population is at risk (Hrabar and Kerley 2015), with one confirmed case in Mountain Zebra National Park (Taplin et al. 2015). If we subtracted the number of individuals (1,346) currently co-occurring Plains Zebra (Hrabar and Kerley 2015) from the total eligible population, an estimated 1,641–2,237 pure individuals remain. Repeating the same calculation including all individuals have previously been kept with Plains Zebra (2,959 in total), yields a mature population size of 753–1,027 pure individuals. Although there is currently no evidence that hybridisation has occurred or that their relative abundance threatens the genetic integrity of Cape Mountain Zebra subpopulations (sensu Piett et al. 2015), these calculations highlight the need to systematically test existing Cape Mountain Zebra populations for genetic purity to more accurately estimate total mature population size.
The population size in 2009 on formally protected areas alone was 1,032–1,408 mature individuals, and 1,385–1,889 in total, which satisfies the IUCN rule of not meeting a threat category for at least five years (IUCN Standards and Petitions Subcomittee 2014).
An intensive survey of Cape Mountain Zebra numbers in 2009 revealed that there were at least 2,790 animals on both formally protected and private properties in the Western and Eastern Cape (Hrabar and Kerley 2013). These surveys were based on aerial surveys for national parks and questionnaires for private landowners and thus the estimated population size is based on reasonably robust data. The survey revealed that, of the 52 subpopulations (compared to 29 in 2004), 17 were formally protected (1,888 individuals) and 35 were privately owned (902 individuals). The survey has recently been repeated (in 2014/15) and it was found that the total population has grown to over 4,790 animals in 75 subpopulations (Hrabar and Kerley 2015). Fifty six populations (1,487 individuals) are on privately owned land and 19 are on formally protected areas (3,304 individuals). The majority of the population (69%) remains on formally protected land and the proportion on privately-owned land (31%) has not risen since 2009, despite the increase in subpopulation number. The MZNP and Karoo NP subpopulations continue to make up a significant proportion of the population, namely 25% and 18%, respectively. Interestingly, Karoo NP’s contribution to the population has remained stable, at 18%, since 2002 and the proportion on MZNP shows a tendency to increase from 20% in 2002 to 22% in 2009 and now 25% in 2015. If it is argued that Cape Mountain Conservation relies on the existence of MZNP, removing the subpopulation leaves 1,927 individuals in formally protected areas remaining, which, at worst, equates to 1,060 mature individuals (55% mature structure. However, when adding the eligible private subpopulations, this increases population size to 3,139 individuals, which corresponds to a minimum of 1,726 mature individuals. Even in the absence of MZNP, there is no continuing decline as other protected areas exhibit an average annual growth rate 8.3% from 2009-2014 (a five year period).
The increase in available suitable habitat, is one reason responsible for the sustained growth rate: for example, Anysberg and Gamkaberg nature reserves have both been expanded in area and a number of stewardship agreements (contract nature reserve) are underway – some of which specifically favour the establishment of Cape Mountain Zebra. Privately owned land played a crucial role in the conservation of the Cape Mountain Zebra when the last few groups in the Cradock area were saved from extinction by local farmers in the 1930s (Skead 2011). This subpopulation was formally protected in 1937 by the proclamation of the MZNP, which was expanded in 1964 to incorporate Cape Mountain Zebra groups occurring on neighbouring private farms (Penzhorn 1975). The expansion of formally protected areas such as the MZNP and Karoo National Park have allowed the growth of the two largest subpopulations. The subsequent increase of the MZNP subpopulation enabled the translocation of individuals to 25 other protected areas during the 1980s and early 1990s, a number of which were private game ranches (Novellie et al. 2002). Similarly, Eastern Cape Parks and Tourism Agency (and its predecessors) have (since 2002) removed 235 Cape Mountain Zebra from Commando Drift and Tsolwana Nature Reserves (166 from Command Drift and 69 from Tsolwana), of which 29 were translocated to the Baviaanskloof Nature Reserve and a further 206 were sold to the private sector. The translocation of animals out of established subpopulations not only reduces density dependent feedback in these subpopulations but creates subpopulations in new areas, each with the potential to increase, while at the same time securing additional habitat. This approach has greatly improved the conservation status of the Cape Mountain Zebra. Subpopulations maintained by private landowners have since increased considerably (Hrabar and Kerley 2015).
Demographic data from nine subpopulations from the Western Cape (in systems with minimal mortality/few predators) reveal a mature population size of ca 67% based on average numbers of mature individuals in both breeding and stallion herds (C. Birss unpubl. data): To compensate for variation between areas, we use a mature population structure of 55–75%. Similarly generation length has been calculated as 16 years (C. Birss unpub. data): The age of first reproduction for females is five years and three months, and remain fertile for 21 years; females first foal at ca 5 years and 3 months; inter-foal periods are 25 months (0.5 foals / year; 12 month gestation period); individuals live up to ca 26 years; and there is ca 26% mortality in foals (Lloyd and Rasa 1989). This is higher than the 11 years estimated for Equus zebra overall by Pacifici et al. (2013). The average breeding group size ranges from 3.4–3.8 individuals (Klingel 1968, Penzhorn 1984, Smith et al. 2008). Bachelor group size has been estimated at 2.5±1 (Lloyd and Rasa 1989). Generation length has been calculated as 10.4 and 8.6 years for males and females, respectively, based on data from De Hoop Nature Reserve from 1995-1999 (Smith et al. 2008). Similarly, Smith et al. (2008) calculate that, to maintain an effective subpopulation size of 50 individuals (thus preventing a significant loss of genetic diversity), 78 individuals should be present at the end of the breeding season (based on ten males breeding annually). A theoretical minimum subpopulation would be composed of ten herd stallions, seventeen bachelor males, 24 females of breeding age and 27 immature animals. Effective subpopulation size will change as subpopulation parameters change (Smith et al. 2008). For example, on the basis of the performance of different subpopulations, Novellie et al. (1996) suggested a minimum founder number of 14 individuals. By only including subpopulations of above 50 (N = 17 subpopulations) and 78 (N = 11 subpopulations) individuals, the total mature population size is estimated to range between 1,758–2,691 individuals.