Cheetah (Acinonyx jubatus)- Data, Pictures & Videos

[Acinonyx sp.]

Cheetah with bay topi kill

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[Acinonyx sp.]

Early Post-Release Movements, Prey Preference and Habitat Selection of Reintroduced Cheetah (Acinonyx jubatus) in Liwonde National Park, Malawi

Focal Species: The Cheetah (Acinonyx jubatus)

2.1. Distribution
 Historically widespread throughout Africa and southwestern Asia, cheetah (Acinonyx jubatus) have disappeared from the majority of their historical range (Durant et al., 2017). Thirty-three remnant cheetah populations are now scattered across 32 of their 53 range states, comprising 9 % of their historical distribution (Durant et al., 2015, Durant et al., 2017). In Asia, cheetah distribution is now limited to the deserts of Iran with an estimated population size of 40, while African population strongholds remain in eastern and southern Africa with an estimated 2290 and 4297 individuals, respectively (Durant et al., 2017). In southern Africa cheetah populations occur across Namibia, Botswana, South Africa and Zimbabwe with remnant populations in Mozambique, Zambia and Angola (Durant et al., 2017; Purchase et al., 2007). During the 2007 and 2017 population assessments, Malawi was the only southern African range state where cheetah were extirpated (Durant et al., 2017; Purchase et al., 2007). Cheetah were present in Malawi in 1975, with an estimated population of 50 individuals spread across two national parks (Gros, 1996). However, by 1989 this population was confined to Kasungu National Park and believed to be mainly transient with neighbouring Zambia (Gros, 1996; Myers, 1975). Recommendations were made to maintain a protected corridor between Zambia’s Luangwa Valley and Malawi to allow for the re-colonization of cheetah in Malawi’s northwestern parks (Gros, 1996). Unfortunately, increased human population growth resulted in the decrease of habitat and prey base within Malawi’s protected areas (PAs) and the unprotected corridor with Zambia (Purchase & Purchase, 2007). The reduction of habitat and prey base, along with the depletion of the Luangwa Valley cheetah population is believed to have prevented any population rescue events in Malawi, resulting in the full extirpation of cheetah from Malawi by the early 1990s (Gros, 1996; Purchase & Purchase, 2007).

2.2. Morphology

The cheetah is one of Africa’s iconic carnivores due to its extreme speed, reaching up to 110 km/h within a few seconds (Marker & Dickman, 2003). This impressive morphological ability is a trade-off for overall bulk, made clear in its anatomy when compared to other large carnivores. The cheetah has an aerodynamically efficient frame with long foot and leg bones that allow for increased speed, and their eyes are positioned for maximum binocular vision and nostrils are enlarged to increase oxygen flow (Ewer, 1973). There are currently five debated sub-species of cheetah (Durant et al., 2017). However, few thorough genetic or morphological comparisons have been conducted. Genetic analysis that has occurred on subspecies has shown a recent divergence (4400-6100 years ago) between the African and Asian subspecies (O’Brien et al., 2017). Morphological measurements appear to vary regionally, which has been attributed to resource availability (Boast et al., 2013; Marker & Dickman, 2003), and while sexual dimorphism is present in the cheetah, it also remains understudied. Therefore, body mass ranges from 35-65 kg and height ranges from 70-90 cm (Estes, 2012).

2.3. Reproduction and cub survival
 Female cheetah become reproductively active around 26-36 months of age (Bissett & Bernard, 2011; Kelly et al., 1998; Laurenson, Caro & Borner, 1992; Marker et al., 2003) and have a litter of one to eight cubs after a 90 – 95 day gestation period (Bissett & Bernard, 2011; Estes, 2012). Cubs are altricial and are born in a den where they remain for 51-65 days (Laurenson, 1993). Upon emergence from the den, cubs remain dependent on their mothers until 14-20 months of age, at which point they separate and form an adolescent group (Bissett & Bernard, 2011; Laurenson, 1993; Marker et al., 2003). An early study examining genetic viability in the cheetah hypothesized that genetic monomorphism resulted in breeding difficulties and high juvenile mortality (O’Brien et al., 1985). Laurenson, Wielebnowski and Caro (1995) later disputed these assertions citing data from three captive breeding institutions in which 78.8 % of cub mortality was attributed to extrinsic factors and 3.8 % to congenital defects. While the paucity of overall genome variability poses conservation challenges for the cheetah, it is unlikely rate-limiting; otherwise the species would not have expanded in range and population size following their genetic bottleneck, which occurred 10-thousand years ago (Menotti-Raymond & O’Brien, 1993; O’Brien et al., 2017). Many cases of breeding depressions in captive cheetah have since been attributed to the species susceptibility to asymmetric reproductive ageing, which, moderates reproductive performance and further reduces genetic variability in captive populations (Ludwig et al., 2019; Wachter et al., 2011). Cub survival in wild cheetah populations appears to be mostly moderated by predation. High occurrences of predation on cheetah cubs were first documented by Laurenson et al. (1992) in the Serengeti National Park, where only 27.7 % of litters emerged from dens. Predation by large predators, mainly lion (Panthera leo) was attributed to 73.2 % of cub mortalities (Laurenson, 1994). Predation continues to be deemed an important cause of cub mortality; however, the rate of occurrences appears to vary across the species range. In the Kgalagadi Transfrontier Park, predation was an important factor of cub survival but not rate limiting, with 35.7 % of cubs reaching adolescence (Mills & Mills, 2014) as opposed to 4.8 % in the Serengeti (Laurenson, 1994). In South African small fenced reserves with varying densities of large carnivores, cub survival post-emergence was 60.0 % (Bissett & Bernard, 2011), and on Phinda Resource Reserve, South Africa, where the lion population was low, cub survival post den emergence was 75.0 % (Hunter, 1998). In Namibia, in areas absent of spotted hyena (Crocuta crocuta) and lion, studies found that between 50.0-78.5 % of cubs monitored from emergence to independence survived (Marker et al., 2003; Wachter et al., 2011). In comparison, leopard (Panthera pardus) cub survival rate in an open system such as the Sabi Sand Game Reserve, South Africa, was 37.0 % (Balme et al., 2012) and lion cub survival was 86.9 % on South African small fenced reserves where infanticide is uncommon (Miller & Funston, 2014). The similarities between cheetah cub survival rates outside of the Serengeti National Park and those of other large African felids demonstrates that the 4.8 % survival rate recorded in the Serengeti National Park may be an outlier due to extrinsic factors. It has been suggested that vegetative cover is important for concealment and thus can influence cheetah cub survival (Durant, 1998; Mills & Mills, 2014). This hypothesis was tested in the Masai Mara, Kenya, by Broekhuis (2018) who determined that habitat density and tourism abundance affected cub survival regardless of lion and hyena abundance. Therefore, habitat homogeneity and elevated levels of tourism in the Serengeti National Park study site may contribute to low cub survival. This highlights the importance of habitat heterogeneity and tourism quotas, before large predator control, when considering habitat protection and reintroduction programs for cheetah conservation (Broekhuis, 2018). 


2.4. Sociality and territorially
 A facultatively social carnivore, the cheetah can form three different social groupings, namely; mother and cubs, adolescent cubs, and male coalitions (Bissett & Bernard, 2007; Durant, Kelly & Caro, 2002; Eaton, 1968). Although adult female cheetah have been found on multiple occasions resting and travelling together, these groupings are poorly understood and believed to be temporary (Dalton et al., 2013). Male coalitions are the most complex social grouping for cheetah, as they have been shown to consist of both related and nonrelated males (Caro & Collins, 1987). Male coalitions are the only long-term social grouping and are attributed to greater territory control, which increases both survival and female encounter rates when compared to nomadic males (Bradshaw, 2014; Caro & Collins, 1987; Caro, Fitzgibbon & Holts, 1989). Home range size varies based on a multitude of factors, including prey distribution, human conflict, patch suitability, distribution of females and presence of other large carnivores (Bissett & Bernard, 2007; Broomhall, Mills & du Toit, 2003; Houser, Somers & Boast, 2009; Hunter, 1998; Purchase & du Toit, 2000; Marker et al., 2007; Rostro-Garcia, Kamler & Hunter, 2015). Cheetah home ranges have been found to span from 32 km2 on small fenced reserves in South Africa to 1651 km2 on Namibian farmland (Bissett & Bernard, 2007; Marker et al., 2007). Changes or disturbances in social grouping also influence home range sizes and can induce shifts. For example, home ranges have been found to increase when a male coalition is reduced to a single male (Marker et al., 2007), whereas female home range greatly reduces during denning (Houser et al., 2009). Despite variation in home range size, the area of core utilization (50 % home range) appears to remain relatively consistent between 10.0 - 13.9 % of the overall home range size (Broomhall et al., 2003; Houser et al., 2009; Marker et al., 2007; Marnewick & Somers, 2015). While home range overlap between male-female ranges is attributed to breeding, male-male overlap and female-female overlap has been shown to also occur at higher than expected frequencies (Broomhall et al., 2003; Marker et al., 2007; Welch et al., 2015). The small area of core utilization may, therefore, account for the high frequency of overlapping home ranges.


2.5. Habitat selection 
Due to its complexity, habitat selection should be viewed as a scale sensitive process and analysed as such (Mitchell & Hebblewhite, 2012). Most commonly examined as a nested hierarchy, habitat selection has been described in four orders; geographic range (1st order), location of home ranges (2nd order), use of habitat within the home range (3rd order) and selection of foraging sites (4th order; Johnston, 1980). Habitat selection should further be considered as a time-sensitive process as seasonality can affect selection in most species (Mitchell & Hebblewhite, 2012). Factors found to affect habitat selection in cheetah include kleptoparasitism, anthropogenic pressures, presence of conspecifics, and prey abundance (Durant, 1998; Klaassen & Broekhuis, 2018; Rostro-Garcia et al., 2015; Welch et al., 2015). These elements are dynamic and can affect habitat selection at different intensities for each selection order. The historical distribution of the cheetah spans most of Africa and into Asia, thereby encompassing multiple biomes. However, early studies of the behavioural ecology of the cheetah centred on the plains of the Serengeti National Park and erroneously deemed the cheetah a grassland specialist (see, Caro & Collins, 1987; Durant, 1998; Durant et al., 1988; Fitzgibbon, 1990; Kelly et al., 1998; Schaller, 1968). Improvement in and the decreasing cost of tracking technology however, increased research capabilities and demonstrated that the cheetah can successfully exploit a wide range of woodland, thicket and arid habitats (Bissett & Bernard, 2007; Broomhall et al., 2003; Cristescu, Bernard & Krause, 2013; Klaassen & Broekhuis, 2018; Marker et al., 2007; Mills, Broomhall & du Toit, 2004; Nghikembua et al., 2016; Rostro-Garcia et al., 2015; Welch et al., 2015). The behavioural flexibility of the cheetah is now evident at the home range scale (2nd order) when comparing selection drivers across study sites. For example, anthropogenic activity and abundance of competing large carnivores appears to have the greatest effect on the location of home ranges for cheetah in open systems (Durant, 1998; Klaassen & Broekhuis, 2018; Van der Weyde et al., 2017) whereas, prey abundance is a driver in home range selection for cheetah fenced systems where they are unable to escape competition (Broomhall et al., 2003; Rostro-Garcia et al., 2015; Welch et al., 2015). Factors affecting 3rd order selection, selection of habitat within the home range, also varies at a study site level. For example, cheetah in Matusadona National Park, Zimbabwe selected open grasslands for hunting and wooded areas for resting and travelling (Purchase & du Toit, 2000). Whereas in South African reserves such as Kwandwe Game Reserve (Bissett & Bernard, 2007), Phinda Private Resource Reserve (Rostro-Garcia et al., 2015), Mountain Zebra National Park (Welch et al., 2015), and Kruger National Park (Broomhall et al., 2003) habitat selection varied based on sex, with females selecting for thicket vegetation significantly more than males. The 4th order of habitat selection for cheetah has not been as thoroughly investigated as that of 3rd order selection. However, semi-closed habitats appear to be selected as kill sites when available, irrespective of prey density, and this is likely a response to kleptoparasitism (Rostro-Garcia et al., 2015). This is further supported by the fact that kleptoparasitism decreases across study sites as cover increases (Mills et al., 2004). Therefore, environmental features related to cover are important in habitat selection of the cheetah on a smaller scale as it is used for the spatial avoidance of intraguild competitors. Anthropogenic factors and prey abundance, however, are thought to affect habitat selection at a larger scale. 

2.6. Prey preference
 Cheetah require less food per day than other large African carnivores (Lindsey et al., 2011). Captive cheetah are fed an average of 1.3 kg/day to maintain a healthy condition (Dierenfeld, 1993). However, calorie needs for captive animals are often lower than those in the wild due to a more sedentary lifestyle. Regardless, wild cheetah consumption rates have been found to range from 0.4 kg/day (Mills et al., 2004) to 4.0 kg/day (Schaller, 1968). Discrepancies in consumption rates are affected by levels of kleptoparasitism, the consumable biomass of captured prey and competition between conspecifics such as cubs or coalition members at a kill (Mills et al., 2004; Schaller, 1968). Due to the morphological limitations of the cheetah, the size range of catchable prey is reduced compared to that of other large carnivores (Hayward et al., 2006). The upper limits of prey that a lone cheetah can successfully capture while minimizing the risk of injury is estimated at 56 kg, while the optimal prey mass has been defined as approximately 27 kg (Hayward et al., 2006). Studies on the prey preferences of cheetah reflect these size limitations as well as demonstrate effects of local prey abundances on preference, with the most abundant medium-sized prey being preferred; impala (Aepyceros melampus) in Matusadona National Park (Purchase & du Toit, 2000), Kruger National Park (Mills et al., 2004) and the Northern Tuli Game Reserve (Craig, Brassine & Parker, 2017), and Thomson’s gazelles (Eudorcas thomsonii) in the Serengeti (Schaller, 1968). However, on Kwandwe Game Reserve in South Africa, greater kudu (Tragelaphus strepsiceros) were the preferred prey species regardless of their large size (>120 kg) (Bissett & Bernard, 2007). The preference towards kudu on Kwandwe Game Reserve is thought to reflect how prey preferences can be altered based on small-scale preferences when the studied population does not mimic natural composition. Small-scale influences on prey preference are therefore important to consider when analysing prey requirements for the desired cheetah population during reintroductions (Lindsey et al., 2011). Male cheetah have been found to hunt larger prey than females in certain populations (Bissett & Bernard, 2007; Mills et al., 2004; Tambling et al., 2014). However, in conflicting studies, no significant differences between male and female prey preferences were found (Clements, Tambling & Kerley, 2016). Contradictory reports in sex disparity for prey preference are attributed to the social dynamics at the study site level, whereas studies in which prey preferences varied greatly between male and female cheetah, often consisted of multiple male coalitions rather than lone males. This indicates that group structure rather than sexual dimorphism is modifying perceived prey preference (Clements et al., 2016). The higher nutritional demands of cheetah social groups are thought to rationalize the preference of larger prey items. While male coalitions have been documented hunting cooperatively (Bissett & Bernard, 2007), females with dependent cubs have been seen to increase their kill rate to account for the additional nutritional demands (Schaller, 1968) further emphasizing the cheetah’s predatory limitations.The presence of intraguild competition and the composition of prey populations have also been shown to affect prey preference. While it has been predicted that areas with low levels of kleptoparasitism would result in cheetah consuming larger prey (Bissett & Bernard, 2007; Hayward et al., 2006), varying densities of competing predators have not been found to influence the size of prey chosen (Clements et al., 2016). Rather, high levels of kleptoparasitism could reflect denser habitat type selected for hunting, and prey preference a factor of predator avoidance behaviours reducing or altering prey options (Clements et al., 2016; Hayward et al., 2006). Finally, cheetah have been shown to display a preference for male antelopes (Fitzgibbon, 1990). This preference is attributed to the reduced vigilance and solitary behaviour of male antelopes as well as the frequency in which they are found on the periphery of groups (Fitzgibbon, 1990; Mills et al., 2004).


2.7. Intraguild competition
 Cheetah are described as a subordinate carnivore as they are reported to suffer from intraguild competition with spotted hyena, lion, and leopard (Hunter, Durant & Caro, 2007; Rostro-Garcia et al., 2015). In the Serengeti National Park, intraguild predation accounts for 73.4 % of cheetah cub mortalities (Laurenson, 1994; Laurenson et al., 1992). While in Matusadona National Park, intraguild predation along with kleptoparasitism is believed to have caused the cheetah population to remain well below the estimated carrying capacity (40 cheetah), with only a maximum of 17 cheetah recorded in the population (Purchase & du Toit, 2000). This demonstrates the significance of interspecific interactions on population dynamics within the large African carnivore community. Cheetah show predator avoidance behaviour by seeking spatial and/or temporal refuges from dominant carnivores (Durant, 1998; Rostro-Garcia et al., 2015). Factors affecting which predator avoidance strategies are exhibited have been correlated to the size of suitable habitat, densities of dominant carnivores and prey availability (Bissett & Bernard, 2007; Durant, 1998; Rostro-Garcia et al., 2015). In Namibia and Botswana, cheetah show large-scale spatial avoidance of predators by inhabiting farmlands where lion and leopard populations are low, as opposed to PAs where dominate carnivore populations are large (Klein, 2006; Marker & Dickman, 2004). However, cheetah on fenced reserves must adopt different predator avoidance strategies. In small fenced PAs cheetah commonly demonstrate temporal avoidance behaviours as there is little room for spatial avoidance, this has been documented by Bissett & Bernard (2007) on Kwandwe Game Reserve (160 km2 ) and by Cristescu et al. (2013) on Shamwari Game Reserve (250 km2 ). In both studies cheetah home ranges overlapped with that of the lion. Avoidance was therefore demonstrated in the form of activity pattern variation as well as habitat preference where cheetah selected for denser habitats, which were commonly avoided by lion. In larger PAs habitat selection appears to be the common mechanism used for minimizing intraguild interactions (Mills et al., 2004; Purchase & du Toit, 2000; Rostro-Garcia et al., 2015). Habitat selection in the form of predator avoidance is a trade-off between resource acquisition and intraguild competition (RostroGarcia et al., 2015). Sex disparities are revealed in this trade-off as cubs are at the greatest risk of predation;therefore females have been shown to utilize thicker habitats at a greater frequency than males (Bissett & Bernard, 2007; Mills et al., 2004; Rostro-Garcia et al., 2015). In the Serengeti, selection of dense habitats is not as feasible, here cheetah demonstrate avoidance and decreased hunting attempts when lion or hyena vocalizations are heard (Durant, 2000a), as well as exhibit multiple prey handling strategies to reduce kleptoparasitism and intraguild predation (Hilborn et al., 2018). This demonstrates that predator avoidance is a flexible behaviour strategy that is adapted based on an individual’s sex, habitat availability as well as prey and predator densities. Furthermore, predator avoidance behaviour in cheetah has been identified as a learned behaviour that is reinforced by breeding success (Durant, 2000b) and predator naïve cheetah are thought to have lower reintroduction success (V. Van der Merwe, pers. comm.).

2.8 Conservation status
 Currently listed as Vulnerable by the IUCN Red-List, it is the large-ranging nature of the cheetah that has been recognized as the main cause for its global population decline (Durant et al., 2017). Once present throughout Africa and into Asia, the cheetah population is now fragmented into 33 populations across 32 range states, totalling an estimated 6700 to 7100 individuals (Durant et al., 2017; Durant et al., 2015). The current range for cheetah consists of only 9 % of their historical range which is primarily found outside of protected areas (Durant et al., 2017). Scenario modelling has demonstrated that cheetah are dependent on PAs as populations outside of these areas are suppressed (Durant et al., 2017). Therefore, population growth inside PAs must remain high to compensate for declines outside of these areas (Durant et al., 2017). Consequently, cheetah are extremely susceptible to a loss of habitat and prey base, as habitat conversion continues in-line with human population growth throughout their remaining range (Durant et al., 2017; Houser et al., 2009; Klein, 2006; Purchase et al., 2007). Currently, over half of the world’s cheetah population presides in six counties within southern Africa; Angola, Botswana, Mozambique, Namibia, South Africa and Zambia (Durant et al., 2017). Cheetah conservation initiatives are broad and differ regionally based on socio-economic and locality factors. In South Africa, a managed metapopulation was developed from cheetah relocated from farmland as part of a conflict resolution initiative (Buk et al., 2018; Lindsey et al., 2011). The metapopulation assists in maintaining a genetically viable population on fenced reserves and PAs in South Africa (Buk et al., 2018). Cheetah often seek spatial refuge from competing carnivores, the practice of fencing of South African reserves and PAs, therefore, prevents cheetah from moving onto farmlands and requires the individual to seek refuge on a small scale (Cristescu et al., 2013). Fenced reserves have therefore been found to maintain carnivore populations closer to their estimated carrying capacities than unfenced areas (Lindsey et al., 2011; Minin et al., 2013; Packer et al., 2013). Cheetah are commonly subjected to kleptoparasitism and at times predation by lion, leopard, and spotted hyena (Durant, 2000b; Durant, 1998). Consequently, cheetah populations outside of fenced PAs are at risk of collapse as they fall into ecological traps while seeking refuge from intraguild competition (Marker & Dickman, 2004). This phenomenon has been recorded in free-ranging populations in Namibia and South Africa, which reside outside of PAs and are highly subjected to retaliatory killings (Marker & Dickman, 2004; Marnewick & Somers, 2015; Muntifering et al., 2006). Conservation of these populations requires intense human-wildlife conflict mitigation in order to increase human tolerance of cheetah (Marker & Dickman, 2004; Purchase et al., 2007). The trade in cheetah poses another sizeable risk to remaining wild populations. Cheetah are included in CITES Appendix I with export quotas for live specimens and hunting trophies from Botswana, Namibia and Zimbabwe (Nowell, CAT & IUCN, 2014). The trade in live specimens poses a unique threat to cheetah compared to other large carnivores. Cheetah can habituate relatively easily especially if obtained while young, and this has primarily fuelled the demand for cheetah as pets (Nowell et al., 2014). Whilst east Africa exhibits the highest records of illegal trade; South Africa has the highest number of breeding facilities and legal live specimen exports (Nowell et al., 2014). Only two of South Africa's breeding facilities, however, are CITES accredited and many boast an unusually high breeding success rate when compared to North American captive facilities (Nowell et al., 2014). This success has resulted in concerns that live-trapped wild animals are illegally entering the legal captive export trade (Buk & Marnewick, 2010; Nowell et al., 2014). South Africa’s legal trade in captive cheetah and the illegal trade in east Africa have therefore been flagged as threats to wild populations in southern and eastern Africa by CITES (Nowell et al., 2014). However, South Africa has recently implemented genetic passports for captive cheetah in order to prove parentage. Genetic passports are now a requirement for the export of cheetah from South Africa, thus have closed a legislative gap that allowed for the illicit laundering of wild cheetah under the guise of the captive trade (Selier & Marnewick, 2019). Whilst the increase in trade regulations are promising, the varying threats, conservation requirements, and sizeable knowledge gap for populations outside of intensely studied PAs has resulted in a formal recommendation for the up-listing of cheetah under IUCN Red List criterion A3b to Endangered (Durant et al., 2017).


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[Acinonyx sp.]

Aspects of Cheetah (Acinonyx jubatus) Biology, Ecology and Conservation Strategies on Namibian Farmlands


1.1 The importance of carnivores
 Carnivores are important indicators of functioning ecosystems and, through predation, impact on all aspects of the system, by diverting what they do not need for their own energetic requirements to scavengers, detritivores, and microorganisms (Ricklefs 1990). However, large carnivore populations are declining globally, with 22 of 30 large carnivore species considered endangered (Fuller 1995), and all are subject to a multitude of pressures, including habitat degradation, conflict with agriculture, hunting, disease and commercial trade (Sillero-Zubiri and Laurenson 2001). Of the carnivores, all the 36 species of Felidae are either classified as threatened or endangered, except for the domestic cat (Felis catus) (Nowell and Jackson 1996). Through their evolutionary history, carnivores have helped shape the evolution of their prey by hunting selection, which has provoked the development of fitnessenhancing anti-predator strategies (Logan and Sweanor 2001). In addition, carnivores have influenced human evolution by enhancing our senses against predation, and we may in part owe the evolution of our large brain and reasoning abilities to carnivores (Wilson 1980). Through scavenging predator kills for themselves, early hominids were provided a high-quality food source that may have enabled them to emerge from Africa and inhabit the globe (Blumenschine 1991, Logan and Sweanor 2001). Carnivores are still influencing human lifestyles through predation on livestock, pets and people. However, their beauty, intelligence and cryptic behaviour has garnered our human curiosity to investigate carnivore species’ evolution, biology and ecology and the consequences for them of living in today’s human dominated landscape. Felids, in particular, are intriguing, and not only provide us with sources of companionship but have provided us with knowledge on genetic links to hereditary defects and diseases that affect both humans and cats through the collaboration between the felid genome organisation and the Human Genome Project (Nash and O'Brien 1982, Rettenberger et al. 1995, Wienberg et al. 1997). Through the study of cheetahs in the early 1980s, today we possibly know more about this species than most other cat species (Caro 1994, Nowell and Jackson 1996, O'Brien et al. 1985, O'Brien et al. 1983), and yet its survival is still in question. In this introduction I summarise information on the cheetah and review its current status, including reasons for its decline. I introduce the background and aims of the research conducted for this thesis, and summarise the contents of each chapter. 


1.2 Cheetah evolution
 An evolutionary history of the cheetah has been constructed by paleontologists from fossils and, more recently, by geneticists using DNA (Adams 1979, Driscoll et al. 2002, Johnson and O'Brien 1997, Menotti-Raymond and O'Brien 1993, van Valkenburgh et al. 1990). Present records date carnivores to the Eocene epoch (Vaughan 1997), about fifty million years ago, with the specialised family Felidae evolving in the Miocene about twenty million years ago. In the middle Miocene, early felids began their radiation into other cats with conical canines including the early cheetahs, Miracinonyx and Acinonyx, during the Pliocene and Pleistocene epochs, about eight million to twelve thousand years ago (Hunt 1996). The cheetah is considered one of the earliest divergences in felid evolution, about 8.5 million years ago, compared to the large cats of the Panthera group, which still shared a common ancestor about 6 million years ago (Adams 1979, Hemmer 1978, Johnson and O'Brien 1997, Neff 1983, Pecon-Slattery and O'Brien 1998, van Valkenburgh et al. 1990). The species known as Acinonyx pardinensis (Adams 1979), which is larger than the modern species, migrated from North America to Asia, India, Europe, and Africa. The modern cheetah evolved into its present form about 200,000 years ago. Genetic research has shown that today’s cheetah populations are descendants of but a few animals that remained after the Pleistocene era about 10,000 years ago, at which point the population experienced a founder event generally referred to as a population bottleneck (Menotti-Raymond and O'Brien 1993, O'Brien et al. 1985, O'Brien et al. 1983). The cheetah somehow survived this time of mass extinction and the population gradually increased. The cheetah was first classified as Felis jubatus (Schreber 1776), but early taxonomists soon realised that the cheetah was unique from all the other cats and placed it into the monospecific genus Acinonyx Brooks (1828), of which there is only the one species jubatus. The translation of the cheetah’s scientific name Acinonyx jubatus is a reference to the species’ semi-retractile pointed claws. In Greek, a means not, kaina, means a thorn, and onus, means a claw (Gotch 1979). A more direct translation may be non-moving claws, and jubatus, in Latin means maned, as young cheetahs have a crest or mane on the shoulders and back. Although seven subspecies have been identified, five subspecies are considered valid by most taxonomists (Smithers 1975). These are Acinonyx jubatus venaticus (Griffith 1821), Acinonyx jubatus hecki (Hilzheimer 1913), Acinonyx jubatus soemmeringii (Fitzinger 1855), Acinonyx jubatus raineyii (Heller 1913), (Schreber, Chapter One – General Introduction 4 1776) and Acinonyx jubatus raddei (Hilzheimer 1913) and details of their distribution are given in Appendix 1. This thesis concerns Acinonyx jubatus jubatus, which is found in Southern Africa.


1.3 Cheetah anatomy and behaviour 
The cheetah is markedly different in both anatomy and behaviour than the other 35 species of Felidae (Ewer 1973, O'Brien et al. 1983). It is the fastest land mammal over short distances (300-400m) (Gray 1968), and has the optimum body size and stride length to reach these high speeds. Nearing full speed, the cheetah is running at about one stride per 0.28 seconds or 3.5 strides per second (Hildebrand 1959, Hildebrand 1961). Due to the cheetah’s specialisation for speed, it has developed many morphological and physiological adaptations. For aerodynamics, it has a small head, lightweight and thinly-boned skull, flat face, and a reduced length of muzzle that allows the large eyes to be positioned for maximum binocular vision, enlarged nostrils, and extensive air-filled sinuses (Ewer 1973). Its body is narrow and lightweight with long, slender feet and legs and specialised muscles, which act, simultaneously, for high acceleration and allow for greater swing to the limbs (Hildebrand 1959, Hildebrand 1961, Neff 1983). The cheetah is the only cat with short, blunt claws, which lack skin sheaths, making the claws semi-retractable, thus providing added traction like a sprinter’s cleats (Ewer 1973). To facilitate the explosion of energy necessary to reach such high speeds, cheetahs are endowed with a powerful enlarged heart, oversized liver, adrenals, bronchi, lungs and large, strong arteries (Eaton 1974, O'Brien et al. 1983). During its high-speed chase in pursuit of prey, the cheetah’s respiratory rate climbs from 60 to 150 breaths per minute, and its body temperature has been measured at 400 C (1050 F; 40 F higher than normal) during a 375-metre sprint (Chinery 1979). For increased intake of air, the nasal passages have become enlarged, crowding the roots of the cheetah’s canine teeth, thus the reason for their smaller size relative to other felids (Ewer 1973). The distinguishing marks of a cheetah are the long tear-drop shaped lines on each side of the nose from the corner of its eyes to its mouth. The cheetah’s coat is tan to a yellow-buff colour, with smaller, less distinct spots between larger spots, and a white belly. Near the end of the tail, the spots merge to form several dark rings. The tail often ends in a bushy white tuft. Although male cheetah are often slightly bigger than females (Caro 1994, Eaton 1974, Wrogemann 1975) and have slightly larger heads, males and females are difficult to tell apart by appearance alone. Cubs are born fully furred and with black spots on a greyish coat. Within two weeks the cubs eyes are open and the fur on the cub’s back begins to grow; by six weeks old the cubs have a long mantle of tan and black fur. Until recently, the cheetah has generally been considered to be an animal of open country and grasslands. This impression is probably due to the ease of sighting cheetahs in the shorter grass, and the long-term studies conducted on cheetahs in East Africa (Caro 1994, Caro and Laurenson 1994, Schaller 1968). However, cheetahs use a wider variety of habitats and are often found in dense vegetation, e.g. the Kora Reserve in Kenya, Botswana’s Okavango Delta, and Namibian farmlands (Broomhall 2001, Marker-Kraus et al. 1996). Even though it is customised for speed, the cheetah can run only 300 to 400 metres before it is exhausted; at this time the animal is extremely vulnerable to other predators, which may not only steal its prey but attack it as well (Caro 1994). Cheetahs are primarily diurnal, possibly due to the nocturnal behaviour of competing predators (Nowell and Jackson 1996). It has been suggested that the cheetah has larger litter sizes as a strategy to offset high juvenile mortality caused by lions and hyaenas (Burney 1980, Caro 1994, Hamilton 1986, Laurenson et al. 1995). Cheetahs have been observed scavenging and returning to a kill, but this is not common behaviour (Burney 1980, Caro 1982, Graham 1966, Pienaar 1969, Stander 1990). Cheetahs also are known to remain on kills in areas where lions and hyaenas are not present (Nowell and Jackson 1996). Cheetahs are considered more social than most other felids, with the exception of the lion (Caro 1994). Large groups of cheetahs (up to 19 individuals of different age groups) have been observed and reported in Namibia and east Africa (Graham 1966, Marker-Kraus et al. 1996, McVittie 1979). Male and female siblings tend to stay together for several months after independence from their dam (Caro 1994), and male littermates remain together in coalitions (Caro 1994). Males in coalitions have been reported to better hold and defend territories (Caro 1994), were found to be in better physical condition and had better access to females for breeding than solitary males (Caro 1994, Caro and Collins 1987). There is considerable variation in cheetah prey, ranging from Thomson’s gazelle (Gazella thomsoni) on the Serengeti plains (Schaller 1968), impala (Aepyceros melampus) in Kruger National Park (Broomhall 2001, Mills and Biggs 1993, Pienaar 1969) to kudu (Tragelaphus strepsiceros), gerenuk (Litocranius walleri) and dik-dik (Madoqua kirkii) in the arid areas of northern Kenya (Hamilton 1986). Other species reported as prey include puku (Kobus vardoni), kob (Adenota kob) and oribi (Ourebia ourebi) (Nowell and Jackson 1996), springbok (Antidorcas marsupialis) (Mills 1990, Nowell and Jackson 1996, Smithers 1975), wildebeest (Connochaetes taurinus) (Eaton 1974, Skinner and Smithers 1990), hare (Lepus spp.) (Labuschagne 1979), and seasonally a large proportion of prey consumed consists of immature ungulates (Burney 1980, McLaughlin 1970). Although, in the central livestock farmlands of Namibia, kudu, warthog (Phacochoerus aethiopicus), red hartebeest (Alcelaphus buselaphus), gemsbok (Oryz gazella), steenbok (Raphicerus campstris) and duiker (Sylvicapra grimmia) have been noted as regular prey species (Marker-Kraus et al. 1996, Morsbach 1987), there has been no quantification of prey consumed in farmland areas. 

1.4 The cheetah’s early association with humans
 The earliest record of the cheetah’s long association with humans dates back to the Sumerians, 3,000 BC, where a leashed cheetah, with what appears to be a hood on its head, is depicted on an official seal (Grzimek 1972, Guggisberg 1975). It was believed in Egyptian history that the cheetah would quickly carry away the Pharaoh’s spirit to the afterlife (Wrogemann 1975)and symbols of cheetahs have been found on many statues and paintings in royal tombs (Guggisberg 1975). Cheetahs were used for hunting in Libya during the reign of the pharaohs (Harper 1945). Cheetahs were not hunted to obtain food, but for the challenge of sport, known as coursing (Guggisberg 1975, Kingdon 1977). In Italy, cheetahs were coursed during the fifth century (Guggisberg 1975, Harper 1945). Russian princes hunted with cheetahs in the 11th and 12th centuries, and, at the same time, crusaders saw cheetahs being used to hunt gazelles in Syria and Palestine (Grzimek 1972). The best records of cheetahs having been kept by royalty, from Europe to China, are from the 14th, 15th and 16th centuries (Guggisberg 1975). Cheetahs also were used for hunting in Russia (Novikov 1956). Eighteenth and 19th century paintings indicate that the cheetah rivalled dogs in popularity as hunting companions (Wrogemann 1975). During his 49-year reign as an Indian Mogul in the 16th century, Akbar the Great had more than 39,000 cheetahs in total, which were called Khasa or the Imperial Cheetahs, and he kept detailed records of them (Caro 1994, Guggisberg 1975). However, all the cheetahs kept for hunting and coursing purposes were taken out of the wild from free-ranging populations. Because of this continuous drain on the wild populations, the numbers of cheetahs declined throughout Asia. In the early 1900s, India and Iran began to import cheetahs from Africa for hunting purposes (Pocock 1939). In Africa, the cheetah was important to many local ethnic groups: the San hunting communities of southern Africa ate cheetah meat for speed; traditional healers used cheetah foot bones for fleet-footedness; and kings wore cheetah skins for dignity (Nowell and Jackson 1996, Wrogemann 1975). These practices, combined with exportation to other countries, contributed to the beginning of the cheetah’s decline in Africa. 

1.5 Current status of the cheetah and population threats
 The cheetah was once one of the most widely distributed of all land animals (Wrogemann 1975). Through the course of time, the cheetah migrated over land bridges from North America into China, through Asia, India, Europe, and finally to Africa (Adams 1979, Kurten 1968, Kurten and Anderson 1980, Martin et al. 1977, Martin and Bateson 1986, van Valkenburgh et al. 1990), settling in its worldwide range as recently as 20,000 years ago (Adams 1979, Wrogemann 1975).In 1900, approximately 100,000 cheetahs were found in at least 44 countries throughout Africa and Asia (Myers 1975, Figure 1.1). The current free-ranging African populations of cheetahs are found in small, fragmented areas spread in 29 African countries of North Africa, the Sahel, East and southern Africa, and it is estimated that around 15,000 animals remain (Marker 1998, Nowell and Jackson 1996, see Figure 1.1), representing a decline of nearly 90% over the century (Marker 1998, see Appendix 1). However, current information about the status of the cheetah in many countries, especially countries that have been engaged in long civil wars, is lacking (Breitenmoser 1998, Breitenmoser and Breitenmoser 2001, Nowell and Jackson 1996). The information from North and West Africa is particularly limited, and the cheetah’s future in these areas is questionable (Marker 1998, O'Mopsan 1998). The remaining strongholds are Kenya and Tanzania in East Africa, and Namibia, Botswana and Zimbabwe in southern Africa (Marker 1998). Cheetah numbers throughout their ranges are declining due to loss and fragmentation of habitat, and a declining prey base (Nowell and Jackson 1996). Intraguild competition from more aggressive predators decrease cheetah survivability in protected game reserves, causing larger numbers of cheetahs to live outside protected areas and therefore coming into conflict with humans (Caro 1994, Marker 1998, Nowell and Jackson 1996). As human populations change the landscape of Africa by increasing the numbers of livestock and fenced game farms throughout the cheetah’s range, addressing this conflict may become the most important factor in their conservation.A further concern is that cheetahs breed poorly in captivity (Marker 2002) and wild populations have continued to sustain captive ones (Marker 2002, Appendix II). Until the 1960s, most cheetahs were imported from East Africa (Marker-Kraus 1997) but, as the numbers of cheetahs decreased in this region, Namibia became the major exporter of cheetahs (Marker-Kraus 1997). Today more than 90% of all cheetahs in captivity are descendants of Namibian cheetahs (Marker 2000, Marker-Kraus 1997). This additional pressure, together with ineffective captive breeding programmes, further endanger cheetah populations. A potentially critical factor for the long-term persistence of the cheetah is its lack of genetic variation relative to other felids. The genetic structure of the cheetah has received considerable attention over the past several years (Driscoll et al. 2002, May 1995, Menotti-Raymond and O'Brien 1993, Merola 1996, O'Brien et al. 1985, O'Brien et al. 1987, O'Brien et al. 1983). It has been suggested that the genetic homogeneity could make the species more susceptible to ecological and environmental changes (MenottiRaymond and O'Brien 1993, O'Brien et al. 1985, O'Brien et al. 1987, O'Brien et al. 1983). This has been interpreted in the context of two potential risks, including the expression of recessive deleterious alleles, and increased vulnerability to viral and parasitic epizootics that can affect genetically uniform populations (Brown et al. 1993, Evermann et al. 1988, Heeney et al. 1990, Munson et al. 1993, O'Brien et al. 1985). Given the lack of genetic diversity, monitoring the overall health of cheetah populations is an important component of understanding and promoting long-term viability (Munson and Marker-Kraus 1997). Over the past few years, the impact of infectious diseases on endangered species has become well known (Burrows et al. 1994, Munson et al. 1993, Roelke et al. 1993, Roelke-Parker et al. 1996). Cheetahs are known to be very susceptible to several feline diseases, and are possibly more vulnerable to such diseases due to the lack of heterogeneity in the population (Evermann et al. 1988, Munson 1993, Munson et al. 1993, O'Brien et al. 1985). In addition, captive populations world-wide have been known to have a high prevalence of unusual diseases that are rare in other species, and these diseases impede the goal of maintaining self-sustaining populations (Bartels et al. 2001, Munson 1993). Although the specific causes of these diseases are not known, the character of these diseases implicate stress as an important underlying factor, and genetic predisposition and diet are possible confounding factors. While it is assumed that these diseases did not historically affect wild populations, there is concern that these diseases may arise in wild animals that are trapped, held in captive facilities and translocated.Additionally, there is concern that cheetahs may transmit or acquire infectious diseases through these actions. Viable populations may be found in less than half of the countries where cheetahs still exist. All populations are listed on the Convention on International Trade in Endangered Species of Fauna and Flora (CITES) Appendix I and are classified as Vulnerable or Endangered by The World Conservation Union (IUCN) (CITES 1984, CITES 1992). The largest remaining wild population of cheetahs is found in Namibia (Kraus and Marker-Kraus 1991, Marker 1998), and these are the subjects of this thesis. 1.6 The cheetah in Namibia Ninety-five percent of Namibia's cheetahs live on the commercial livestock farmland, which covers 275,000 km2 of the country's north central region (see Figure 1.2) (Marker-Kraus and Kraus 1990, Morsbach 1987). The widespread removal of lions (Panthera leo) and spotted hyaenas (Crocuta crocuta) from commercial farming areas early in the 1900s opened a niche for the cheetah to fill. The abundance of water and natural prey animals on these farms allowed the cheetah to successfully inhabit these areas (Marker-Kraus et al. 1996). Although the cheetah has been reported to be declining in numbers throughout its range (Hamilton 1986, Joubert 1984, Kraus and Marker-Kraus 1992, Marker 1998, Marker-Kraus and Kraus 1990, Myers 1986, Stuart and Wilson 1988, Wilson 1987, Wrogemann 1975), little research has been conducted outside fenced game reserves or protected areas, despite the fact these are now the most important habitats for cheetah in Namibia. At the same time, studies on captive cheetah have yielded extensive information about their biology, physiology, and behaviour (Brown et al. 1993, Dierenfeld 1993, Evermann et al. 1988, Howard et al. 1993, Lindburg et al. 1993, Marker-Kraus and Grisham 1993, Munson et al. 1993, Wildt et al. 1993, Wildt and Grisham 1993), but there have been no comparable studies conducted on free-ranging cheetahs. In southern Africa, cheetahs are killed regularly in farming areas due to their raiding of livestock and the attitudes of the farmers (Marker-Kraus et al. 1996, Marker-Kraus et al. 1993, Morsbach 1987, Stuart and Wilson 1988, Wilson 1987). Although classified as a protected animal in Namibia, a cheetah can be shot in order to protect one's life or property (Marker-Kraus et al. 1996). Between 1980 and 1991, 6,829 cheetahs were legally removed from the wild Namibian population, mainly through indiscriminate catching in live traps and shooting (CITES 1992). Carnivore-livestock conflict has been exacerbated by a change in husbandry during the past century (Breitenmoser 1998). For instance, in recent decades, domestic livestock is no longer herded or guarded by dogs and as such is more vulnerable to predation. Furthermore, stockmen have lost the tradition of coexistence with large predators and modern protective legislation of carnivores is not matched by positive cooperative attitudes by livestock communities (Breitenmoser 1998). The increasing availability of illegal and legal firearms is also likely to pose a threat so long as the cheetah’s skin has any value (Hamilton 1986). 

1.7 Livestock management and non-lethal predator control
 The key question for management of large carnivores in today’s human dominated landscape is what is the impact of predators on livestock? There are a variety of reasons for livestock loss, including disease, poor management, and predation. Because of this, identifying the correct cause of livestock loss is fundamental. Worldwide farmers commonly blame predators for the majority of livestock losses before investigating the cause of loss thoroughly (Marker-Kraus et al. 1996). If predation is discovered as the cause, then identifying the appropriate culprit is necessary for effective management strategies to be undertaken. When the specific predator causing livestock loss has been accurately determined, identifying livestock management strategies to help prevent further losses is then necessary. All information and management practices must be evaluated carefully since every situation is unique, and different methods may be required for reducing predation in each case. Some farmers, through the world indicate that implementing new livestock protection methods is too much work, as their management was already extensive (Landry 1999). This attitude is unfortunate, as the lack of any predator control or the presence of vulnerable livestock or game may encourage opportunities for predators, and improper methods of farming can create losses to predators. Conversely, proper management can prevent or remedy many problems. A large variety of management practices have been used successfully in Namibia to reduce livestock loss to cheetahs (Marker-Kraus et al. 1996). Some of these strategies include calving camps, corralling calving herds and utilising guard animals such as donkeys or baboons that can reduce loss to predation. Predation on cattle calves may decline if farms synchronise calving both within their herd and with other farms in the area, as well as with wildlife calving times. High concentrations of cattle calving within a short time period has helped, as there is protection in numbers. This, combined with a fast rotation schedule through smaller camps, has helped several farmers. Farmers that breed more aggressive breeds of cattle have shown to have lower losses to predators, as these breeds are more protective of their calves. Inexperienced heifers calving for the first time should be given additional protection, such as putting them with older cows or in closely observed calving camps. Calving seasons are critical, especially for heifers. It is best for them to calve in mid-summer when there are more wild young, as well as more cows and calves for protection, as the first calves born during the start of a calving season are the most likely to be killed. A cow that fails to reproduce or loses its calf to predation should be culled from the herd. The use of donkeys to protect livestock from predators has been used in many areas of the world (Landry 1999, Marker-Kraus et al. 1996). Donkeys are generally docile, but are know to have an inherent dislike for intruders such as cheetahs, blackbacked jackals, caracals and even domestic dogs. Farmer benefits include low cost, easy management, and a high success rate (Marker-Kraus et al. 1996). Mules also have been used for protection because they are more aggressive than donkeys. Although mules are aggressive guard animals, they have been known to ‘steal’ calves for themselves, since they cannot reproduce. Zebras, horse stallions and horned oxen have been used successfully to deter predators. The early Namibian settlers commonly kept horned oxen with their calving herds (Marker-Kraus et al. 1996). Farmers have also expressed the belief that cattle, especially females, should never be dehorned; and that mature cattle are more successful against predators than heifers (Marker-Kraus et al. 1996). A few farmers have used baboons to protect smallstock, however the aggressive behaviour of the baboons eventually prevents even the farmers from getting near the flock (MarkerKraus et al. 1996). Other forms of predator control included poison collars on stock to selectively eliminate the specific livestock-killing animal; and both sight and sound repellents, which can be effective temporary aids to protect livestock, but predators soon become accustomed to the repellents (Landry 1999). In addition, taste aversion has been experimented with on canids, causing the predator physical illness after eating treated bait (Gustavson et al. 1976). This method has proved very effective, as it targets the specific problem animal. Smallstock farmers often use dogs with their herds. The majority of the dogs used in Namibia to protect livestock showed ‘herding dog’ behaviour instead of the appropriate ‘guarding dog’ behaviour (Marker-Kraus et al. 1996). Herding dogs use the eye-stalk behaviour to move stock, similar to predators. It is believed that when a predator approaches the herd, the dog instinctively begins to herd the animals, due to the herding instinct. This stimulates the predatory motor pattern of the predator (eye-stalkchase-trip-bite-consume), causing it to chase and kill the stock. Conversely, specialised breeds of ‘livestock guarding dogs’ discourage the predatory behaviour, as guarding dogs act as sentries. When a predator approaches, a guarding dog will bark while moving towards the predator, and then will retreat back into the herd, without causing the herd to run. This pattern is repeated, thus breaking the predatory motor pattern. Predators usually are opportunistic and will seek prey elsewhere once challenged. This may be especially true of the non-aggressive cheetah. Therefore, using a selected breed of livestock guarding dog could be an effective non-lethal predator management strategy for farmers in Namibia. 

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[Acinonyx sp.]

Mask of blood: Cheetahs' faces are covered in gore after feasting on a kill

• The cheetahs were coated in crimson after they devoured a 30lb antelope in Kenya's Maasai Mara reserve 
• Vivid images show the so-called coalition of cheetahs licking their lips and prowling the game reserve
• Pictures of the 'thrilling moment' were taken by 41-year-old photographer Tapan Sheth from Rajkot, India 
  

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[Acinonyx sp.]

Cheetahs feeding on wildebeest in Kwandwe great fish river lodge

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[Acinonyx sp.]

Cheetahs return to Angola's south


Wild cheetahs have returned to southern Angola for the first time in decades, having disappeared during decades of civil war, a researcher said Friday.

I was in southern Angola to make a survey, looking for signs of cheetahs, and we were just ecstatic to find cheetahs there," said Laurie Marker, from the Cheetah Conservation Fund in neighbouring Namibia.
"I actually saw two wild cheetahs, which is very rare, to visibly see them," she said.
"To be able to see wildlife starting to come back is a huge benefit for Angola and it is wonderful news at a biodiversity level in general," she added.
The cats were seen in the Iona region in southern Namibe province, home to Angola's biggest national park, which was badly damaged during the 27-year civil war that ended in 2002.
Angola's environment ministry in January declared 2010 the "year of biodiversity", saying it wanted to restore its parks and create new conservation areas.
Marker said southern Angola could develop eco-tourism, which is the backbone of the tourist trade around the region.
"There is a great potential for tourism. But they have to be very cautious and delicate" not to destroy this wildlife that is only slowly returning.
Last year researchers discovered a rare Angolan antelope that had been feared extinct, spotting three of the giant black sable that are a national symbol.
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[Acinonyx sp.]

There have been less than 200-500 cheetahs in Angola in the 1970's

From The Cheetah Acinonyx Jubatus in Africa


http://www.catsg.org/cheetah/05_library/...Africa.pdf

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[Acinonyx sp.]

Cheetahs in Kidepo

Cheetahs are rare animals that can hardly be seen in any national park but In Uganda they are found in the pearl of Africa hidden treasure in the most remote national park that can't easily be accessed either.


The park is situated in the north-eastern part of the country.Cheetahs belong to the cat family and can only be seen in Kidepo national park in Uganda.Some years ago there were a lot of fears that cheetahs may have been extinct in kidepo valley national park but reports have it that they are spotted in park though not oftenly.


There were reports from the Uganda wildlife that there are only about 20 cheetahs in kidepo valley national park.Therefore something must be done to protect them and their habitat to avoid more extinction there are several non-governmental organisations implementing conservation programs to reduce the poaching activities and over use of animal habitats the best time for viewing cheetahs and other big cats is before sunrise and at sunsets


However apart from the rare cheetahs,the world's largest bird can also be sighted in Kidepo in large groups-the ostriches not to be recorded in any other national park in Uganda.

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[Cheetah9750]

Cheetah winks

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[Acinonyx sp.]

Power output of skinned skeletal muscle fibres from the cheetah (Acinonyx jubatus)

SUMMARY

Muscle samples were taken from the gluteus, semitendinosus and longissimus muscles of a captive cheetah immediately after euthanasia. Fibres were ‘skinned’ to remove all membranes, leaving the contractile filament array intact and functional. Segments of skinned fibres from these cheetah muscles and from rabbit psoas muscle were activated at 20°C by a temperature-jump protocol. Step and ramp length changes were imposed after active stress had developed. The stiffness of the non-contractile ends of the fibres (series elastic component) was measured at two different stress values in each fibre; stiffness was strongly dependent on stress. Using these stiffness values, the speed of shortening of the contractile component was evaluated, and hence the power it was producing. Fibres were analysed for myosin heavy chain content using gel electrophoresis, and identified as either slow (type I) or fast (type II). The power output of cheetah type II fibre segments was 92.5±4.3 W kg−1 (mean ± s.e., 14 fibres) during shortening at relative stress 0.15 (the stress during shortening/isometric stress). For rabbit psoas fibre segments (presumably type IIX) the corresponding value was significantly higher (P<0.001), 119.7±6.2 W kg−1 (mean ± s.e., 7 fibres). These values are our best estimates of the maximum power output under the conditions used here. Thus, the contractile filament power from cheetah was less than that of rabbit when maximally activated at 20°C, and does not account for the superior locomotor performance of the cheetah.

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[Acinonyx sp.]

Sexual dimorphism in cheetahs

4.3.1 Sexual dimorphism 
Significant sexual dimorphism, with males being larger, was evident for all measurements recorded, and is summarized in Table 4.2. There was a significant difference between the mean weights of male cheetahs between years (F = 2.880, d.f. = 8, P = 0.007) but not for females (F = 1.136, d.f. = 8, P = 0.368)

Table 4.2 Morphometric data for the wild adult cheetahs (aged over 30 m at capture, and held in captivity for <30 days), collected using the protocol shown in Appendix IV. All measurements are in centimetres (cm), apart from body mass, which is measured in kilograms (kg). Statistical tests were conducted between the measurements for adult males and females to ascertain the degree of sexual dimorphism exhibited for each parameter. *Test 1 = independent samples t-test, with equal variances assumed (test statistic = t), and test 2 = Mann-Whitney U test (test statistic = z). 

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[Acinonyx sp.]

Cheetah Cubs starting to eat meat

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[Acinonyx sp.]

LYMPHOSARCOMA ASSOCIATED WITH FELINE LEUKAEMIA VIRUS INFECTION IN A CAPTIVE NAMIBIAN CHEETAH 

ABSTRACT 
This case report describes the occurrence of multicentric lymphosarcoma in a four- year-old female wild-born captive-held cheetah in Namibia after being housed in an enclosure adjacent to a feline leukaemia virus (FeLV) infected cheetah that had previously been in contact with domestic cats. The year prior to the onset of clinical signs, the wild-born cheetah was FeLV antigen negative. The cheetah subsequently developed lymphosarcoma, was then found to have seroconverted to FeLV, and then rapidly deteriorated and died. At necropsy, the liver, spleen, lymph nodes and multiple other organs were extensively infiltrated with neoplastic T-lymphocytes. FeLV DNA was identified in neoplastic lymphocytes from multiple organs by PCR and Southern blot analysis. Although the outcome of infection in this cheetah resembles that of FeLV infections in domestic cats, the transmission across an enclosure fence was unusual and may indicate a heightened susceptibility to infection in cheetahs. Caution should be exercised in holding and translocating cheetahs where contact could be made with FeLV-infected domestic or feral felids. 

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[Cheetah9750]

Khatu is a rewilded cheetah and also a super mom, living successfully on her own at Buffalo Kloof private reserve in South Africa and had birth to six little healthy cubs.

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[Acinonyx sp.]

CHAPTER 8: THE INCIDENCE OF DENTAL ABNORMALITIES IN WILD CAUGHT NAMIBIAN CHEETAHS 

ABSTRACT 
Two hundred and eight cheetahs that were opportunistically live-trapped on Namibian farmlands were examined for signs of dental abnormalities. Three abnormalities were recorded: erosion of the upper palate, possibly a predisposition to focal palatine erosion, where the first lower molar penetrates the palatine mucosa; crowding of the lower incisors; and the absence of one or both upper premolars. Just over 40% of the cheetahs examined showed deep palatine erosion, and 15.3% of these had perforated upper palates. In addition, 31.7% of the cheetahs examined had crowded lower incisors and 20.9% had one or both upper premolars missing. The incidence of focal palatine erosion is of particular interest as it has only been recorded in captive cheetahs, where it was attributed to a ‘soft’ captive diet, and never before in completely wild individuals. To attempt further understanding of the potential causes of such erosion, the condition was examined in relation to sex, age, region, time in captivity, and the occurrence of other dental abnormalities. No relationship was found between the severity of the condition and time spent in captivity, while juveniles showed more severe erosion than adult cheetahs. Cheetahs missing either one or both upper premolars showed a higher incidence of deep erosion, and the same was true for cheetahs that exhibited crowded lower incisors. The traditional explanation of focal palatine erosion being an artifact of captivity does not explain its occurrence in this sample population of cheetahs, the majority of which were raised entirely in the wild.

8.1 INTRODUCTION
 Focal palatine erosion is a health problem that has been identified in captive cheetahs, in which the first lower molar penetrates the upper palate, medial to the upper first molar (Fitch and Fagan 1982, Phillips et al. 1993). In unaffected cheetahs, there is a slight indentation of the palatine mucosa in this general area, to accommodate the cusp of the tip of the lower first molar. This erosion is a serious problem that has the potential to cause severe health problems and even fatal disease in individuals, especially when penetration of the hard palate occurs. The pathogenesis of FPE occurs where the lower molar’s tip makes regular contact with the palatine mucosa, so the tooth eventually penetrates through the palatine bone itself, causing inflammation. The oral defects observed in affected cheetahs range from sparse cellulitis, the loss of pigmentation and signs of inflammation, to large oral-nasal bony defects extending through the palatine bone into the nasal passage (Fitch and Fagan 1982). Particles of food which lodge in the focal palatine defect result in localised infection and further tissue damage. FPE has been reported in captive cheetahs as early as 10 months of age with a slight, localised cellulitis, although in young cats it may be overlooked as a typical ‘teething’ disorder (Fitch and Fagan 1982).Although this condition has never previously been reported in cheetahs that were born and raised in the wild, the majority of reported cases have occurred in Namibian wild-caught animals living in captivity and captive-born animals from Namibian founders. When first reported, 86% of the cheetahs with FPE came from one shipment from Namibia in 1970 or their descendants (Fitch and Fagan 1982). This oral defect was attributed to the feeding of soft commercial diets lacking bones on developing cheetahs (Phillips et al. 1993), as well as the possibility of specific family lines, renal disease, suppurative rhinitis, and appears to often accompany, but not always, a maloccluded dentition (Fitch and Fagan 1982). During examinations of cheetahs handled by the Cheetah Conservation Fund, the opportunity was taken to investigate any dental abnormalities that were observed in wild cheetahs. In addition to focal palatine erosion, other dental abnormalities observed in our sample population were the crowding of the lower incisors, and the absence of one or both upper premolars. The crowding varies from slight crookedness to a severe condition where the incisors are arranged in two parallel rows. In domestic dogs and cats, such problems usually have a genetic basis, although nutritional status, juvenile viral infections, and metabolic disorders are also possible causes (Colmery and Frost 1986, Frost and Williams 1986). The absence of one or more premolars has been recorded in cheetahs before (Ewer 1973), but the objective in this study was to examine this phenomenon in relation to the other abnormalities recorded, particularly the erosion of the upper palate. Before investigation into the impact, prevalence, and etiology of such dental abnormalities can be undertaken, the anomalies must be properly defined, characterized, and described in literature. The reporting of these conditions, particularly focal palatine erosion, in entirely wild cheetahs is important for other researchers, to encourage further investigation, and to aid in the determination of the ultimate causes of focal palatine erosion and its impact on wild cheetahs. 

8.2 METHODS 
Cheetahs were examined after being opportunistically live-trapped on Namibian farms, as described in Chapter 3, and had been held in captive situations for varying lengths of time before the Cheetah Conservation Fund was invited to examine them. The region and date of capture was determined whenever possible, and cheetahs that had been held in captivity for 30 days or more by the time of examination were considered to be ‘captive’ animals. Age classification followed the protocol described in section 3.1, but in order to enable comparisons to be made, we condensed our age classes into the scheme et out in Phillips et al. in (1993), which used four classes for analyses: (1) Juvenile cheetahs (24 months old or less at the time of exam) that had been captured before adult tooth eruption (before seven months old); (2) Juvenile cheetahs captured at seven months old or later; (3) Adult cheetahs (over 24 months old) that had been captured when juvenile (seven to 24 months old) and; (4) Adult cheetahs that were captured over 24 months old. Each cheetah was examined for signs focal palatine erosion (FPE) and evidence of other dental abnormalities. A score of 1 was assigned in cases where there was very little or no sign of erosion, a score of 2 indicated a medium erosion, and in cases where erosion had caused a deep depression the condition was scored as a 3. Some of the cases scored as 3 also showed actual focal palatine erosion, signified by perforation of the mucosa, sometimes accompanied by bleeding, inflammation, and signs of foreign matter, and this was recorded as well. Callipers were used to measure particular teeth and intraoral photographs were taken. The overall score for analysis was developed as follows: 1 = a score of 1 on both sides of the palate 2 = a score of 1 on one side and 2 on the other 3 = a score of 2 on both sides 4 = a score of 2 on one side and 3 on the other 5 = a score of 3 on both sides. None of the cheetahs examined had scores that did not fit this scheme, e.g. a score of 1 on one side and 3 on the other. The degree of erosion was considered to be severe if ne or both sides of the palate were scored with a 3. The number of upper premolars was examined for each cheetah, with resulting scores of 0 (no premolar on either side), 1 (one premolar present on one side) and 2 (both premolars present), while the lower incisors were also examined to see whether there was any crowding. Cheetahs were also weighed and their physical condition assessed, looking at factors such as coat condition, musculature, ectoparasite load etc. This excluded injuries that were likely to have been sustained while in the capture cage, to give a better indication of condition in the wild. Statistical analyses were performed using SPSS version 10.0 software (SPSS Inc. Chicago, USA). Means significance testing was carried out using the parametric independent samples t-test, preceded by Levene’s test for equality of variances, and general linear model univariate analyses. Departures from expected ratios were analyzed using Pearson’s chi-squared test, while the non-parametric Spearman’s rank correlation coefficient was used to determine the significance of relationships between variables measured on ordinal scales. All tests are two-tailed unless otherwise stated. 

8.3 RESULTS 
Two hundred and eight cheetahs were examined for dental abnormalities between June 1992 and November 1999, and the breakdown of this sample population is shown in Table 8.1. Almost two-thirds (62.5%) of the cheetahs examined were male, while a similar proportion (67.3%) were wild, i.e. had been held captive for under 30 days. 

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Precise dates of capture were available for 94.2% (n = 196) of the examined cheetahs and these animals could therefore be assigned to the four age classes described above. Overall, 87 juvenile cheetahs were examined, 12.6% of which (n = 11) were captured before seven months old (Table 8.1). Although the single most common score assigned was 1, the incidence of deep erosion was relatively high, with 40.9% of the sample population classified as having a deep depression on at least one side of the palate (Table 8.2). Thirteen (15.3%) of these cheetahs (6.3% of the sample population) had perforations in the upper palate as a result of this severe focal palatine erosion.

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The frequency of deep erosion amongst cheetahs that had been held captive for 30 days or more was 42.6% (n = 29), while the condition was slightly milder in wild 
cheetahs, with 40.0% (n = 56) showing severe erosion. There were no significant differences between wild and captive cheetahs, however, regarding overall score (F 2 = 3.573, d.f. = 4, P = 0.467), the incidence of severe erosion (F 2 = 0.133, d.f. = 1, P = 0.716), or the development of palatine perforation (F 2 = 1.142, d.f. = 1, P = 0.285). When just captive cheetahs were examined, there was a slight positive correlation between the score assigned and the length of time spent in captivity, although it was not statistically significant (rs = 0.203, n = 57, P = 0.129). Figure 8.1 shows the mean scores by sex for the different age classes. There was no difference between sexes in overall scores (F 2 = 4.980, d.f. = 4, P = 0.289), the frequency of severe erosion (F 2 = 2.017, d.f. = 1, P = 0.156), or perforated FPE (F 2 = 0.005, d.f. = 1, P = 0.941). There was some slight variation, however, in scores between different regions of the country (F = 2.026, d.f. = 8, P = 0.045).

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Age did appear to play a significant role in the severity of erosion: juvenile cheetahs had significantly higher overall scores than the adults (F 2 = 13.645, d.f. = 4, P = 0.009), were subject to significantly more severe erosion (F 2 = 9.930, d.f. = 1, P = 0.002), and were significantly more likely to show perforated erosion (F 2 = 5.971, d.f. = 1, P = 0.015). There was no significant difference in the severity of the erosion seen between those juveniles captured before adult tooth eruption and those captured when older (F 2 = 0.859, d.f. = 1, P = 0.354). Adult cheetahs captured as juveniles had a lower incidence of severe erosion than those captured when adult already, although the difference was not statistically significant (F 2 = 3.725, d.f. = 1, P = 0.054). Data regarding the number of premolars and the frequency of crowded incisors are shown in Table 8.3 for captive and wild cheetahs examined of different ages. 

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The number of premolars that a cheetah had was significantly linked to the severity of erosion – those cheetahs that had either one or both upper premolars missing showed a significantly higher frequency of severe erosion than those with both premolars present (F 2 = 7.251, d.f. = 2, P = 0.027). The frequency of severe erosion was also significantly higher amongst cheetahs that had crowded lower incisors than those that did not (F 2 = 4.537, d.f. = 1, P = 0.033), although there was no significant relationship for perforated FPE (F2 = 0.290, d.f. = 1, P = 0.590). There was a relationship between the incidence of severe erosion in wild cheetahs and poorer physical condition, as shown in Figure 8.2. Wild cheetahs with severe erosion were significantly less likely to be in excellent condition (F 2 = 11.296, d.f. = 1, P < 0.001). While wild cheetahs that had severe erosion were lighter in mass than those without a severe condition, with a mean mass of 33kg compared to 36kg, this difference was not statistically significant (t = 1.203, d.f. = 135, P = 0.231). There was no significant relationship between crowded incisors and either being in excellent physical condition (F 2 = 0.673, d.f. = 1, P = 0.412), or body mass (t = 1.429, d.f. = 135, P = 0.155).

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8.4 DISCUSSION 
Although focal palatine erosion was first formally recorded in the captive cheetah population almost 20 years ago, it was believed not to occur in wild cheetahs captured as adults (Fitch and Fagan 1982, Phillips et al. 1993). During this long-term study conducted on Namibian farmlands, however, we found not only evidence of erosion in wild cheetahs that may lead to focal palatine erosion, but also found that it was not uncommon, with 40% of wild cheetahs showing deep erosion on at least one side of the palate. Even more notably, six percent of the cheetahs examined were suffering from perforated palates (the most severe form of focal palatine erosion), including seven animals that had never been held in captivity. This shows that the preconditions for FPE exist in the free-ranging cheetah population throughout Namibia, with slight variations by region. A common explanation for the development of focal palatine erosion has been that cheetahs brought into a captive situation are fed artificially ‘soft’ diets, which fail to sufficiently wear down the carnassials and hence lead to palate damage (Fitch and Fagan 1982, Phillips et al. 1993). This does not explain, however, the results presented here, which show that cheetahs reared on an entirely natural, wild diet also suffer from the same problem. The effect of diet on tooth wear may indeed be important, and is supported in this study by the evidence that the degree of erosion appears to increase with time spent in captivity, although the trend here was not found to be statistically significant. The fact that the captive cheetahs we examined showed a slightly higher incidence of severe erosion also hints towards some effect of captivity, but we feel that focusing entirely on the issues of diet and captivity as the sole explanation may be misleading. The juvenile cheetahs examined showed a significantly higher degree of erosion than did the adults, regardless of how old they were when they were captured, and were also more likely to suffer from perforated focal palatine erosion. Although wild cheetahs consume primarily muscle and skin (van Valkenburgh 1996), bone consumption at kills has been recorded to varying extents (Brain 1981, Phillips 1993). It seems likely that the gradual wear from gnawing on tough cartilage and bones will eventually blunt the teeth of adult cheetahs and reduce the extent to which the molar can irritate and penetrate the palatine surface. In 1982, Fitch and Fagan suggested that malocclusion of the teeth could be a anomalies similar to those reported here have been observed in the highly inbred populations of captive white tigers (Emily, P., pers. comm.), while other morphological abnormalities have been reported in Florida panthers, which also show very little genetic variability (Johnson et al. 2001, Roelke et al. 1985). The combined allozyme study on Namibian cheetahs and the morphological data suggests a genetic explanation (O'Brien et al. 1983, Wayne et al. 1986). The dental abnormalities as described in this paper may prove to be a genetic condition that predisposes an individual to developing advanced focal palatine erosion. Extensive work has already been done on cheetah genetics (Menotti-Raymond and O'Brien 1993, O'Brien et al. 1985, O'Brien et al. 1987, O'Brien et al. 1983) and protocols exist for both DNA fingerprinting and microsatellite analysis (Gilbert et al. 1991, Menotti-Raymond and O'Brien 1993). Both of these techniques will be used in the near future to examine relatedness of animals with and without these morphological abnormalities. While the occurrence of focal palatine erosion in wild cheetahs is interesting from a scientific standpoint, the most crucial factor is whether it appears to have a detrimental effect on the cheetahs that exhibit it and on the population overall. Phillips et al. (1993) found with captive animals that even those cheetahs that had FPE were in excellent condition, but we found that in our sample there was a relationship between severe FPE and a loss of physical condition. It is not yet known in what way the dental abnormalities described will impact on the cheetah’s ecology, nor what the consequences will be in the long-term. For example, wild cheetahs are known to suffer from kleptoparasitism by larger, more powerful carnivores such as lions and hyaenas (Caro 1994), and if cheetahs with dental abnormalities are slower at processing their kills, they may be at a further disadvantage in the wild by losing more of their kills in this way. It is also unknown what the implications of the physical defects, such as perforated palates as seen in extreme cases of FPE, may be for the overall health of affected wild individuals, although we know that they can lead to serious physical problems in captivity. For instance, one of the captive cheetahs reported with FPE died from severe kidney failure that was associated with oral-nasal osteomyelitis from FPE (Fitch and Fagan 1982), and kidney failure is one of the main causes of death for captive cheetahs (Marker-Kraus 1997, Munson 1993). Overall, the development of dental abnormalities, including focal palatine erosion, is likely to be multifactorial, with genetics, diet, time spent in captivity, age and skull morphology all playing a role. The reporting of these conditions in wild cheetahs is important, as raising awareness amongst researchers will be vital in order to gain more information regarding the prevalence and severity of such abnormalities in different cheetah populations. This information will be crucial for a better understanding of how these conditions, particularly focal palatine erosion, may develop amongst wild cheetahs as well as those in captivity. 

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