Cheetah (Acinonyx jubatus)- Data, Pictures & Videos

[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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