The status of rhinoceroses in South African National Parks

Rhinoceroses (rhinos) are under threat worldwide. Of the African rhinos, the southern white rhino Ceratotherium simum simum is the most abundant (Amin et al. 2006a), with its total numbers exceeding that of a minimum viable population (Reed et al. 2003). In contrast, the African black rhino sub-species experienced severe declines with the eastern black rhino Diceros bicornis michaeli and south-western black rhino Diceros bicornis bicornis listed as critically endangered and the south-central black rhino Diceros bicornis minor listed as vulnerable (Amin et al. 2006a). The northern white rhino Ceretatherium simum cottoni is functionally extinct (Groves, Fernando & Robovský 2010) and the western black rhino Diceros bicornis longipes (Emslie 2011) recently went extinct (Travers, Waterland & Stroud 2011). Asian rhinos (greater one-horn rhino Rhinoceros unicornis; Javan rhino Rhinoceros sondaicus and Sumatran rhino Dicerorhinus sumatrensis) all survive only in small numbers, with the Sumatran and Javan rhinos each with less than 100 in the wild. There were 3557 greater one-horn rhino by the end of 2015 (Emslie et al. 2016).


Introduction
Rhinoceroses (rhinos) are under threat worldwide.Of the African rhinos, the southern white rhino Ceratotherium simum simum is the most abundant (Amin et al. 2006a), with its total numbers exceeding that of a minimum viable population (Reed et al. 2003).In contrast, the African black rhino sub-species experienced severe declines with the eastern black rhino Diceros bicornis michaeli and south-western black rhino Diceros bicornis bicornis listed as critically endangered and the south-central black rhino Diceros bicornis minor listed as vulnerable (Amin et al. 2006a).The northern white rhino Ceretatherium simum cottoni is functionally extinct (Groves, Fernando & Robovský 2010) and the western black rhino Diceros bicornis longipes (Emslie 2011) recently went extinct (Travers, Waterland & Stroud 2011).Asian rhinos (greater one-horn rhino Rhinoceros unicornis; Javan rhino Rhinoceros sondaicus and Sumatran rhino Dicerorhinus sumatrensis) all survive only in small numbers, with the Sumatran and Javan rhinos each with less than 100 in the wild.There were 3557 greater one-horn rhino by the end of 2015 (Emslie et al. 2016).
By 2014, South Africa was home to approximately 90% of the global population of southern white rhinos, and 36% of the world's black rhinos, consisting of south-western, south-central and a small extra-limital population of eastern black rhinos (Emslie et al. 2016).By then, the South African National Parks Board (SANParks) was custodian to roughly half of the white rhinos (49%) and 31% of the black rhinos within South Africa (Emslie et al. 2016).South African national parks are thus home to a substantial proportion of the world's rhino and play a key role in ensuring their survival.
African rhinoceroses (rhinos) experienced a poaching onslaught since 2008 with the epicentre in South Africa where most of the world's rhinos occur.South African national parks, under the management of South African National Parks (SANParks), are custodian to 49% of South Africa's white and 31% of the country's black rhinos.We collated information on rhino population sizes in seven national parks from 2011 to 2015.We include and report on rhino surveys in Kruger National Park during 2014 and 2015.Southwestern black rhinos increased over the study period, which allows SANParks to achieve its contribution to South Africa's 2020 target of 260 individuals.South-central black rhinos declined over the study period because of poaching in the Kruger National Park, making it difficult for SANParks to realise a 9% increase per annum for its expected contribution to the South African target of 2800 individuals.For southern white rhinos, SANParks requires 5% annual growth for its contribution to the South African target of 20 400 individuals.To continue to evaluate the achievement of these targets, SANParks needs annual population estimates relying on total counts, mark-recapture techniques and block-based sample counts to track trends in rhino populations.SANParks' primary challenge in achieving its contribution to South Africa's rhino conservation targets is associated with curbing poaching in Kruger National Park.

Conservation implications:
The status and trends of rhino species in SANParks highlight key challenges associated with achieving the national targets of South Africa.Conservation managers will need to improve the protection of southern white rhino, while the Department of Environmental Affairs need to be made aware of the challenges specifically associated with not achieving targets for south-central black rhino.Outcomes for south-western black rhino have already realised and the good conservation efforts should continue.

The status of rhinoceroses in South African
National Parks

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Several regional, national and international rhino conservation strategies and targets thus underpins SANParks' rhino management.The Black Rhino Biodiversity Management Plan (Knight et al. 2013)  Although authorities recognise the effect of rhino poaching on southern white rhinos in Kruger National Park (Ferreira et al. 2015), there was no previous published attempt to quantify SANParks' overall progress towards the national rhino conservation targets across the SANParks estate.Moreover, SANParks have not yet assessed whether the techniques employed to count rhinos are optimal at current population sizes or densities.This study therefore aims to collate all SANParks' data on rhino population estimates for the period 2011-2015, across the range of methodologies employed to (1) evaluate the population status of southern white rhino, south-western and south-central black rhino populations in SANParks, (2) assess whether the techniques employed to count rhinos are optimal within park-specific contexts and (3) recommend optimal survey designs for the three sub-species in various National Parks.For south-central black and southern white rhinos in Kruger National Park we also derive estimates of recruitment rates (i.e.proportion of rhinos less than 1 year old reflect rhinos born and that survived the first year after birth) between two surveys and compare these to estimates of poaching rates.The results will allow us to evaluate SANParks' progress in achieving South Africa's rhino conservation targets and help inform management interventions.

Study populations, data collection and collation
The study populations include all rhinos in SANParks' protected areas from 2011 to 2015 (Appendix 1).Seven  Ferreira and Greaver (2016).This study made use of a mark-recapture approach to estimate population size.
For Kruger National Park, we collated south-central black rhino data from Ferreira, Greaver and Knight (2011) as well as Ferreira et al. (2015).These publications provided estimates with 95% confidence intervals for 2009 and 2013, respectively.
To obtain estimates for 2011 and 2012, we used Monte-Carlo simulation techniques (Fishman 1995) to randomly draw from the probability distributions of the estimates collated for 2009 and 2013, and calculated an annual exponential growth rate (Caughley 1977).
Our study also made use of surveys targeting the southern parts of Kruger National Park during September 2014 and 2015 using block count techniques (Ferreira et al. 2011(Ferreira et al. , 2015)).
The technique focused on surveying 470 and 487 randomly placed blocks 3 km × 3 km in size during 2014 and 2015, respectively.Surveyors systematically completed transects comprising a 200 m observation strip on each side of a helicopter within each block with flights 45 m above ground at 65 knots.The survey team comprised a pilot, a data recorder and two observers.Using the block count data with an estimator (Jolly 1969) during 2014 and 2015 allowed landscape-specific estimates and overall estimates for Kruger National Park after accounting for the following biases: availability bias came from relationships between vegetation cover and rhino visibility (Ferreira et al. 2015), and observer bias came from estimates made during a previous survey (Ferreira et al. 2011).Detectability bias was minimal as the size of observation strips was narrower than it was in previous studies when detectability was noted as negligible (Kruger, Reilly & Whyte 2008).
Using the observed age distribution and sex ratios, we could derive recruitment rates for Kruger National Park between the surveys of 2013 and 2014, as well as 2014 and 2015.This allowed us to compare the number of rhinos recruited through birth and first year survival in the total black rhino population with the number of rhinos poached between the two surveys.We thus collated poaching data in Kruger National Park for those periods (SANParks Unpublished records 2 ).

Analysis of trends in population estimates and growth rates
We calculated sub-species-specific exponential growth rates for each park using an exponential model (Caughley 1977).
For parks and species that included sample-based estimates with associated 95% confidence intervals in any particular year we made use of Monte Carlo simulations (Fishman 1995) to define exponential population growth rates and their associated 95% confidence intervals.For each year when a sample-based estimate was available, we extracted a value of the population size from the probability distribution defined by the estimate and its 95% confidence interval of that specific population estimate, and calculated exponential growth.We repeated this process 100 000 times and calculated the median, as well as 2.5% and 97.5% percentiles as a definition 2.Data available from Ken Maggs, ken.maggs@sanparks.org3.Data available from Cathy Greaver, cathy.greaver@sanparks.org and Charlene Bissett, charlene.bissett@sanparks.org of the 95% confidence interval.If the confidence intervals of exponential growth rates excluded zero, we concluded that a population is changing.This allowed us to identify the National Parks that play an important role in SANParks' ability to achieve contributions to national rhino objectives.
In order to evaluate SANParks' requirements to meet national targets by 2020, we calculated the annual rhino population growth rate required for each sub-species to reach the stipulated targets, given the population status during 2015 (see the 'Results' section).These evaluations allowed us to assess whether authorities require revised conservation interventions.

Determining optimal survey requirements
Surveying populations living at low densities present similar statistical challenges as surveying populations with few individuals.We generated a two-way matrix that describes the population sizes and densities of all rhino species in SANParks (Table 2).For ease of classification we defined population density categories on exponential scales.This allowed us to place a population in the smallest category based on size or density and identify the most suitable techniques given the category (Table 3).
As the survey techniques employed to estimate southcentral black and southern white rhinos had estimates of confidence intervals, and thus precision (i.e. the likely spread of estimates given the uncertainties introduced by biases such as availability, observer and detectability biases, see Caughley 1974 andThompson 1992), these values allowed us to define optimal monitoring intervals directed at detecting required population growth.We used the required growth rates to achieve targets along with recorded confidence intervals to define optimal survey requirements using power analyses to detect trends (Gerrodette 1987).When designing optimal survey requirements, authorities need to trade-off the magnitude of change to detect, survey intervals, the number of surveys required and the total change by the time a trend is detected.This results in authorities deciding how many surveys and intervals are needed to obtain reliable data for detecting trends.5).

Contribution of various subpopulations to rhino population size targets
Between the 2014 and 2015 surveys, the number of rhinos born and surviving the first year were similar to that poached.This result confirms that, despite significant numbers of southern white rhino poached, Kruger National Park remains the most important contributor towards the conservation of wild free-ranging individuals of this species within its natural range in South Africa.
, Note that techniques such as camera trapping can assist with observations particularly in areas with high levels of vegetation cover.

Contemporary trends and requirements to meet conservation targets
South-western black rhinos increased at a rate of 0.17 (95% CI: 0.10-0.

Optimal survey intervals and requirements
Survey methods for south-western black rhinos (Table 4) do not allow for estimates of confidence intervals and hence do not allow definition of optimal surveys through trade-offs (Gerrodette 1987).Consequently, south-western black rhino numbers require estimation on an annual basis making use of total counts as before and mark-recapture techniques in future using ear-notching (Greaver et al. 2014;Ngene et al. 2011) and or genetic marking (Brook et al. 2012).
In contrast, estimates for SANParks' south-central black rhinos made it possible to generate coefficients of variance http://www.koedoe.co.zaOpen Access  (8% during the study period) with which optimal survey intervals could be calculated.Two surveys between 1 and 4 years apart are needed to detect the required annual increase from 2016 to 2020.At this survey interval, there is a trade-off with the total amount of change detected at the time of the second survey's completion.If declining trends continue as at present, a total reduction of 11% south-central black rhinos will be detected during two surveys 1 year apart.In contrast, a total reduction of 44.2% individuals will be detected with surveys every 4 years (e.g.2016-2020).Optimal survey intervals for south-central black rhinos should also be annual.
Techniques should focus on block counts and mark-recapture estimates in Kruger National Park and mark-recapture approaches elsewhere.
Estimates of southern white rhinos living in South African national parks had a coefficient of variance of 5% at the end of 2015.To detect an annual decline of -0.05 as well as the required annual exponential increase of 0.05 from 2016 to 2020, SANParks can estimate southern white rhinos twice with surveys 1-4 years apart.The total change after 1 year, if potential declining trends realise (i.e. 5% decline in numbers), carries less risk than the total change after 4 years (i.e.22.6% decline in numbers).Southern white rhinos thus also require annual estimates.Techniques should focus on block counts in Kruger National Park and total counts elsewhere.

Discussion
SANParks play a key role in the conservation of three subspecies of rhino within free-ranging conditions in South Africa.For south-central black rhinos and southern white rhinos, however, contemporary trends predict that SANParks will not be able to meet contributions to South Africa's rhino population targets if observed annual population growth rates remain the same.
Encouraging is the significant increases noted for southwestern black rhinos living in Addo Elephant, Mountain Zebra, Karoo and Mokala National Parks.The observed annual growth exceeds the physiological capability of southwestern black rhinos -having their first calf at 5-11 years of age, and at best, a calf every 2 years thereafter (Hitchins & Anderson 1983).The high observed annual growth can result from small population effects on accuracy and precision of population estimates (Gerrodette 1987), accentuating estimated vital rates (Akςakaya 2002).High population growth can also result from conservation husbandry seeking to maximise population growth, including skewing sex ratios (Holand et al. 2003) and moving rhinos to maximise the social requirements of individuals (Reid et al. 2007).Over the past 5 years, SANParks moved three south-western black rhinos between four National Parks, placed five under custodianship and introduced an additional seven individuals from elsewhere into National Parks.The overall growth comes from having a female sex skew resulting from the initial introductions and high female calving rate in the initial years after introduction.).The rise in poaching incidents in Namibia (AfRSG Unpublished data 7 ) reduces the likelihood of sourcing south-western black rhinos from there.An encouraging prospect is that SANParks' conservation husbandry approach from 2011 to 2015 resulted in an inflated increase in south-western black rhino numbers within South African national parks.
South-central black rhinos require a 9% increase in numbers per annum to reach the SANParks contribution to South Africa's 2020 target.This exceeds the maximum growth predicted by best survival, the ages at which cows have their first calves (Hitchins & Anderson 1983) and how often thereafter (Ferreira et al. 2011;Hitchins & Anderson 1983).
In addition, SANParks only have two parks where southcentral black rhinos occur within their natural distribution.Conservation husbandry and expanding Marakele National Park (Ferreira & Greaver 2016) should continue, but these practices were not enough to compensate for the losses experienced in Kruger National Park from 2011 to 2015.The poaching onslaught in Kruger National Park is the most 5.Dave Zimmermann, SANParks, dave.zimmermann@sanparks.org 6.African Rhino Specialist Group, Mike Knight, mike.knight@sanparks.org7.African Rhino Specialist Group, Mike Knight, mike.knight@sanparks.orgsignificant threat to SANParks' ability to achieve its southcentral black rhino target by 2020.
The 5% annual increase required for southern white rhino populations is biologically realistic.In fact, the age at which a cow has her first calf, along with birth intervals and survival rates (Owen-Smith 1988) predict >5% potential annual growth rates.For instance, southern white rhinos realised 9% annual growth in Kruger National Park prior to the poaching onslaught that began in 2008 (Ferreira & Okita-Ouma 2012).
SANParks should continue to implement rhino protection in small parks, anti-poaching in Kruger National Park and biological management across all of its national parks.Implementation has borne some results -in the 18 months before the end of 2015, only one rhino was confirmed poached in the small parks (SANParks Unpublished data 8 ).
The number of confirmed rhinos poached in Kruger National Park was one less during 2015 (826) compared to 2014 (827) (SANParks Unpublished data 11 ).This contrasts the pattern from 2008 to 2014 when there was a 32% annual increase in the number of rhinos poached (SANParks Unpublished data 11 ).SANParks anticipate that the full implementation of zone-, technology-and intelligencedriven anti-poaching in Kruger National Park will provide improved control of poaching within the park.If complimented with game-changing interventions, including disrupting organised crime (Haas & Ferreira 2015) and empowering people (Lunstrum 2013) outside the park, authorities can curb rhino poaching.Several ongoing national and international collaborations, as well as new initiatives could aid in ensuring the long-term persistence of rhinos in Kruger National Park.
The results of our analysis suggest three key directives for conserving south-central black rhinos: (1) the existing conservation targets require revision of the 5% annual increase requirements in the Black Rhino Biodiversity Management Plan (Knight et al. 2013).Given the effect of poaching (Ferreira et al. 2015), the targeted numbers of SANParks' contribution will require higher annual growth rates, (2) taking control and curbing the effect of poaching on south-central black rhinos in Kruger National Park requires urgent interventions including a prioritised security response using interdiction patrol and hot pursuit tactics (Haas & Ferreira 2017;Park et al. 2016) informed by predictions of spatial distribution of rhinos (Rachlow, Kie & Berger 1999) extracted from intensive biological monitoring programmes and (3) we recommend that land for use by south-central black rhinos be increased.The northern part of Kruger National Park provides an ideal opportunity for expansion.
Although south-central black rhinos mostly occur in southern Kruger at present, rangers noted several cases of southcentral black rhinos previously using landscapes in northern Kruger (Ferreira et al. 2011).Expansion into the area can only be considered if authorities adequately address the internal and external factors driving rhino poaching in the park.

Conclusion
The trends of two of the three sub-species of rhinos in national parks are of key concern.Poaching, specifically in Kruger National Park, has disrupted the ability of SANParks to achieve its contribution to South Africa's rhino conservation targets for 2020.Achieving targets is feasible for southwestern black rhinos, less so for southern white rhinos, and most challenging for south-central black rhinos.Twenty-seven rhino orphans (24 white rhino and three black rhino) were rescued from 2012 to 2015 during the increased poaching onslaught.Twenty-four of these orphans were still alive by the end of 2015.Even so, rhinos continue to colonise KNP with 77.1% of landscapes having white rhinos present by 2010.Since 2008, the population was influenced by poaching as well as density-and rainfall-dependent responses in birth and death rates.
National Parks -Kruger, Marakele, Mapungubwe and Mokala National Parks.We collated all southern white rhino observation data for these parks from 2011 to 2015.3 Annual and bi-annual helicopter surveys flying at a height of 50 m at a speed of 50-60 knots used 200 m wide transects to systematically cover a Park.This provided total counts of southern white rhinos at Mokala National Park and Marakele National Parks, respectively.Southern white rhinos in Kruger National Park also occur mostly south of the Olifants River.We collated estimates for 2011, 2012 and 2013(Ferreira et al.  2012(Ferreira et al. , 2015)).During September 2014 and September 2015 we made use of the same survey and analytical approaches as used for south-central black rhinos in Kruger National Park.Rangers have a full record of every individual in Mapungubwe National Park based on regular observations.
Caughley 1974authorities, however, face several challenges when counting and detecting trends in rhinos and other large animal population sizes.These relate to the feasibility of techniques at different population sizes and densities (Table1), and sources of bias or error (e.g.Caughley 1974) linked to different methods and habitats.
(Knight et al. 2015rican black rhino population growth rate of 5% per annum, with 2800 south-central and 260 south-western black rhinos by the end of 2020.The White Rhino Biodiversity Management Plan(Knight et al. 2015) also aims for a white rhino population growth rate of 5% per annum, with 20 400 southern white rhinos by the end of 2020 in South Africa.In addition to these rhino Biodiversity Management Plans (BMPs), South Africa's cabinet adopted an integrated four-pronged approach to curb rhino poaching during 2014 (Department of Environmental Affairs 2014).These are: (As rhino conservation targets are set around rhino population sizes and growth rates, it is important to survey populations frequently to detect statistical changes.Estimates and trends allow for evaluating progress towards the targets, and assessing the effectiveness of current management interventions.

TABLE 1 :
Survey techniques, situation suitability and supporting literature.
Addo Elephant National Park makes the largest contribution towards south-western black rhino populations, with the combined numbers in Kuzuko, Nyathi and Colchester-Main Camp Sections comprising a minimum of 120 individuals (Table4).Over half of the south-central black rhinos within SANParks occur in Kruger National Park (Table4).Between the 2013 and 2014 surveys, the number of south-central black rhinos born that survived the first year in Kruger National Park was similar to the number of rhinos poached (Table5).More south-central black rhinos were poached between the 2014 and 2015 surveys than what were born and survived the first year.
The block count of southern white rhinos in Kruger National Park produced an estimate of 8821 (95% CI: 8335-9307) south of the Olifants River during 2015.This figure increased to a total of 8875 (95% CI: 8365-9337) after including field ranger observations north of the Olifants River (Table5).Elsewhere, an additional 252 individuals occur in Mapungubwe, Mokala and Marakele National Parks combined.More southern white rhinos were born and survived the first year than what were poached between the 2013 and 2014 surveys (Table

TABLE 2 :
Summary of rhino populations within South African National Parks in categories of population sizes (number of rhinos) and densities (n.km -2 ).

of Rhinos Density Density very low: Very low density of rhinos Density < 0.01 n.km -2 : Low density of rhinos Density 0.01 n.km -2 - < 0.1 n.km -2 : Medium density of rhinos Density 0.1 n.km -2 -< 1.0 n.km -2 : High density of rhinos Density 1.0+ n.km -2 : Very high density of rhinos
We provide names of the National Parks.Note that Addo Elephant National Park has three different sections where south-western black rhinos occur.We provide categorisation in a two-way matrix based on absolute number of rhinos as well as density of rhinos because parks have very different areas.SWR, southern white rhino; SWBR, south-western black rhino; SCBR, south-central black rhino.http://www.koedoe.co.zaOpenAccess

TABLE 3 :
A summary of techniques associated with evaluating various aspects of African rhino conservation plans.Note that suitable techniques can be informed by the total size of a rhino population and/or the density of rhinos in an area of interest.

TABLE 4 :
Population estimates of various subspecies of rhinos within South African National Parks.We provide 95% confidence intervals for estimates that made use of formal statistical techniques.

TABLE 5 :
A summary of first year recruitment (the number of rhinos born and surviving the first year of life) and poached individuals recorded for south central black rhinos and southern white rhinos in Kruger National Park between the 2013, 2014 and 2015 surveys.

intervals First year recruitment Poached rhinos South-central black rhino
African national parks?Nonetheless, curbing rhino poaching in Kruger National Park remains the highest priority for SANParks' contribution to South Africa's rhino initiatives.

TABLE 1 -
A1: Features of South African national parks where rhinos occur, along with a brief summary of rhino history in each park.Seven D.b.michaeli were introduced into AENP between1961  and  1962 .By 1996this population had increased to 35 individuals.All the D.b.michaeli were subsequently removed and replaced with the South-western black rhino ecotype D.b.bicornis.The reintroduction of D.b.bicornis to the AENP formed part of SANParks overall policy to re-establish only indigenous species/subspecies/ecotypes in national parks.This reintroduction has gone through a number of phase extending over several isolated management units, as the park has expanded.At present there are three major areas that sustain the population: 1.The Darlington-Kuzuko Section, located in the Nama-Karoo biome 597 km 2 in size, is at present divided into two sections: Kuzuko which is a bull camp only and Darlington Dam section (445 km 2) which has a breeding population of black rhino.The population at Darlington Dam was founded in 1999 with the reintroduction of five rhino from Augrabies Falls National Park.The population was reinforced with a bull from Tswalu in 2005, three (one male and two females) from Namibia in 2005, and a bull from Addo Main Camp in 2007.Three bulls have been removed from the population since the founding of the population.A total of 29 individuals have been notched since 2002.Three bulls were relocated to the Kuzuko section in 2005.This bull group has had various removals and introduction since then.2. Nyathi Section (145 km 2 ) is located in the forest-thicket transition.Between 1994 and 1997, eight black rhinos were reintroduced into this area and the population has since been supplemented with two bulls in May 2003 and June 2003, respectively.Since then there has been no further reinforcements of the population and no removals from the population.Fifteen individuals have been notched since 2003.No rhinos have been poached.3. The Colchester-Main Camp Section is unit (287 km 2 ) and comprise mostly of the thicket biome.Between 1998 and 1999 SANParks introduced a founder population of 15 black rhinos, but experienced numerous mortalities relating to disease and post release stress from elephant-rhino conflict.The population has since been reinforced with two rhino from Tswalu Game Reserve in 2000, a Namibian bull in 2007 who died shortly after reintroduction, another Namibian bull in October 2009 and five from Hunter's Moon in May 2012.Only one bull has been removed from the population since 2003.No rhinos have been poached.A total of 16 individuals were notched since 2003.
Vaalbos National Park to Tswalu Game Reserve.In 2005, one bull was removed from Vaalbos National Park.The initial reintroduction to MoNP was six individuals that were relocated from Vaalbos National Park in 2006, however one female died during relocation.Therefore the starter population was five individuals.In April 2007 an additional 11 animals were relocated from KNP to MoNP and in 2014 another 5 animals were relocated from KNP to MoNP.

TABLE 1 -
A1(Continues...): Features of South African national parks where rhinos occur, along with a brief summary of rhino history in each park.MapNP) is located in the Limpopo Province on the South African side of the confluence of the Shashe and Limpopo Rivers.The park is a World Heritage Site and protects significant cultural and historical assets and is located in the Savanna Biome.The Park is 281 km 2 in size comprising of formally declared protected areas and private land managed by SANParks and experience annual rainfall of 350 mm -400 mm.Winters are mild, while summer temperatures may rise to 45 °C.Commiphora-Colophospermum veld dominates the undulating terrain, while the unique tall Lowveld riparian woodland lines the main rivers.SANParks introduced four southern white rhinos to MapNP during 2004.The population has remained small largely because of poor breeding and natural mortality, poaching and little additional introductions.Since 2008 one rhino was poached in MapNP.Marakele National Park Marakele National Park (MaNP) is 639 km 2 and located in the Savannah biome of the North West Province of South Africa.It includes the Marakele Park (PTY) Ltd. managed as a contractual park.Mountains and bottomlands dominate the landscape.The initial reintroduction of D.b.minor to MaNP was three individuals into the Waterval area (two from KNP and one from Hluhluwe-Imfolozi Park) during October 1993.Subsequent to that 25, additional black rhinos were introduced until 2007.Five were moved out of the park between 2003 and 2008 and some internal translocations have also taken place.Mortalities have been attributed to natural causes.Regular notching events until 2013 also ensured that most of the MaNP black rhino population have unique ear markings to facilitate population monitoring.Southern white rhinos were introduced to MaNP initially during 1996.Subsequent to the initial introduction, 93 additional white rhino were introduced from 2001 to 2015 from KNP.During 2003, two white rhinos were moved to Botswana and another five were removed from 2007 to 2013.Kruger National Park Kruger National Park (KNP) is situated in the low-lying Savanna of South Africa and covers 19 485 km² with annual rainfall exceeding 450 mm.Granite and gneiss deposits separated by Karoo sediment combine with wooded Savanna comprising Sclerocarya caffra and Senegalia nigrescens on basalts and mixed Combretum spp.and Senegalia spp. on granites in southern KNP.Colophospermum woodlands dominate the north to create 35 different landscape types across KNP.South-central black rhino went locally extinct and the last one was sighted during 1936.Reintroduction commenced in 1971, with a total of 81 black rhinos introduced by 1990.By 2009 black rhinos were increasing at 6.8% (95 % CI: 0.041-0.098)per annum and have reached a population size of 627 (95% CI: 588-666).South-eastern black rhino had high survival and an estimated inter-calving interval of 2.45 years.Age distributions and population growth suggested higher survival rates for dependent calves and adults compared to sub-adult males and females.The poaching onslaught since 2008 has resulted in a decline in the population.Several removals (six to Liwonde National Park in Malawi, six to Venetia Game Reserve (seven were captured with one capture related mortality), 12 (13 were caught with one capture related mortality) to North Luangwa National Park in Zambia, six to Moremi Game Reserve in Botswana and two orphans were raised and sold to private owners before 2012) can influence population responses.Southern white rhinos were locally extinct by the turn of the 19th century.Authorities introduced 351 white rhinos between 1960 and 1972 sourced from Hluhluwe-iMfolozi Park.By the middle of the 1980s, authorities started to remove rhinos from KNP as donations to other conservation areas and zoological gardens.The population proliferated and by the late 1990s, authorities sold/exchanged and donated annually between 0.1% and 1.6% of the white rhinos to private owners as part of a developing wildlife economy associated with southern white rhinos and to other conservation areas.Between 1995 and 2015, 1465 white rhinos were moved out of KNP.