An independent literature review commissioned by CapeNature

LITERATURE REVIEW OF THE ECOLOGY AND CONTROL OF THE BLACK-BACKED JACKAL AND CARACAL IN SOUTH AFRICA

by

J du P Bothma

BSc & MSc (University of Pretoria) PhD (Texas A & M University) Pr. Sci. Nat

July 2012

Not intended for general reference

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EXECUTIVE SUMMARY

Carnivore ecology 1. Carnivores have been living in Africa for millions of years but the current carnivores have all only existed for five million years or less. These later carnivores have lived alongside early humans for at least the last two million years but it is only after small livestock were domesticated in the last 12 000 or so years that conflict has developed between humans, their small, domesticated livestock and wild carnivores. 2. The primary causes of this conflict are competition for the same space and natural resources. Moreover, although many carnivores are omnivores, they all also eat meat. 3. Carnivores are ecologically adaptable and occupy specific ranges that are large enough to support their maintenance. These ranges are defended against competing individuals of their kind. 4. Many small, domesticated livestock production units have become degraded ecosystems with a scarcity of natural prey for carnivores and such flocks are targeted by the carnivores as food resources at times. Such production of small, domesticated livestock is essentially an agricultural monoculture and has a severe impact on ecosystem productivity and functioning. 5. Young carnivores are highly productive and show rapid population growth. New territorial animals will soon move into any ranges that become vacant as a result of natural mortalities or carnivore control programmes. 6. The only viable way in which to reduce the impact of carnivores on flocks of small, domesticated livestock permanently is to eliminate the carnivore(s) totally on a regional or national level. This is economically impossible and ecologically undesirable to do. 7. The removal of carnivores from an ecosystem will create imbalances and ecosystem collapse which will reduce the productivity of the system whether for wild or for domesticated herbivores. 8. Carnivores are usually opportunistic feeders which will utilize alternative prey resources; including small, domesticated livestock; when survival requires it to do so. Carnivores cannot regulate abundant prey populations but will regulate small prey populations.

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9. Functional ecosystems are productive and natural prey may partially or totally buffer small, domesticated livestock against depredation by carnivores. In such ecosystems carnivore populations form an interrelated web. 10. Mainly some adult, territorial males in a carnivore population will become habitual killers of small, domesticated livestock. Unselective control programmes may do more harm than good, and when they also kill rare or threatened wildlife they become illegal. 11. In a functional ecosystem wild herbivores can maintain substantial production levels despite the presence of carnivores because of the presence of natural defences against predation. Where such defence mechanisms have been lost in small, domesticated livestock flocks they should be compensated for by livestock husbandry practices that are based on ecosystem principles or be redeveloped through selective breeding. 12. Except the lion, spotted hyaena and brown hyaena none of the carnivores in South Africa is sociable. Consequently they cannot attain high densities as they may live in defended ranges alone (caracals) or as mated adult pairs (black-backed jackals). 13. Predation by carnivores on small, domesticated livestock is most severe during the lambing season(s). 14. Carnivores that weigh less than 14,5 kg preferentially kill prey animals that weigh less than them and are more energetically rewarding to hunt. 15. Predation is a vital part of ecosystem functioning and the dynamics of carnivore populations must include broader environmental principles and issues. The lack of predation in an ecosystem can cause serious productivity imbalances which could end in the overall collapse of ecosystem productivity. The occurrence of predation can be beneficial to prey because of competition between different types of carnivore for space and food and could increase the productivity of a habitat to support herbivores. 16. The development of husbandry programmes for small, domesticated livestock on an ecosystem basis should be based on the interaction between three components: humans, carnivores, livestock and other herbivores. Generalizations are invalid and each programme should be based on the circumstances that prevail. 17. Large, functional ecosystems are self-sustaining and develop a natural balance between carnivores and their prey but this becomes disturbed in dysfunctional ecosystems such as on some production units of small, domesticated livestock.

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18. There are many popular perceptions about the nature, scope, causes and remedies for small, domesticated livestock depredation but most of them are not supported by independent scientific studies. Carnivore-prey interactions and carnivore control 1. Predation is economically and ecologically relevant to the production of small, domesticated livestock and is a vital part of ecosystem functioning. 2. Detrimental cascading biodiversity and production effects may follow where carnivores have been exterminated. 3. The average carnivore in Africa is prone to predation and ecological influence from 15 other types of carnivore at some stage of its life. 4. Carnivore-livestock predation management through the control of carnivore numbers has been practised for centuries throughout the world but except for total extermination of the carnivores involved it has failed in all the properly documented cases. 5. The true benefits and costs of the control of carnivores on small, domesticated livestock production units should not only be measured in the number of carnivores that are being killed but rather in proportion to the true costs of the damage or losses and the benefits that are gained. 6. The benefits of taking an holistic ecosystem approach to the husbandry of small, domesticated livestock should be built into financial management and animal husbandry plans. Small adaptations towards holistic livestock management can make substantial differences to the impact of carnivores on livestock and increase the ability of the habitat to support herbivores. 7. The existence of such holistic financial and husbandry plans could become the point of departure for conservation authorities when considering applications for carnivore control permits. Such plans should include data on the long-term trends in carnivore populations, trends in the productivity of the habitat to support herbivores and the removal, if any, of carnivores over time. 8. Subsidisation from government sources to compensate the producers of small, domesticated livestock for losses that were caused by carnivores has not produced positive results in maintaining or increasing livestock production anywhere in the world.

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The value of using scientific nomenclature 1. When dealing with and analysing depredation and carnivore control, the sex, age and type of the carnivore that is involved have to be identified. 2. There are numerous confusing local and regional common names for organisms which lead to misconceptions and erroneous perceptions. 3. The binary system of the taxonomic classification of all the organisms of the world that was created by Linnaeus in the 1750s has cleared up this confusion and such scientific names should be used to identify the source of any depredation on small, domesticated livestock. 4. The binary classification system of Linnaeus eventually classifies all animals into genera and species, and sometimes into subspecies. A genus which only contains one species is considered to be monotypic. The black-backed jackal 1. The black-backed jackal is an opportunistic hunter and scavenger of medium size with an omnivorous diet. It occurs widely in Africa and in most parts of South Africa except the deep forests. 2. Black-backed jackals have existed for some 2,5 million years and are known to have lived alongside early humans some two to one million years ago. The Khoikhoi pastoralists reported that they had experienced depredation by wild carnivores on their recently domesticated small livestock centuries ago. 3. Some 16 common names in Afrikaans are known for the black-backed jackal in South Africa. This creates confusion and misconception in identifying those animals that are really responsible for depredation on small, domesticated livestock. 4. The black-backed jackal shows a preference for open woodlands and grasslands but occurs ubiquitously in South Africa, only being absent in the deep forests. 5. Mated pairs demarcate defended ranges, the size of which depends on habitat quality, the intensity of human disturbance and the abundance of prey. In functional ecosystems these ranges are small with little overlap, exclude other black-backed jackals and limit overpopulation. 6. Territory boundaries can break down temporarily at an abundant food source such as a large carcass and often creates misconceptions about black-backed jackal population densities.

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7. Young and highly productive black-backed jackals disperse up to 135 km away from their natal sites into ranges that have become vacant because of the natural or control-induced mortalities of black-backed jackals and this rapidly lead to population recovery. This limits the effectiveness of control programmes. 8. Black-backed jackals are active by day and night when they are not being persecuted by humans but they become mainly nocturnally active when being persecuted. 9. When the male member of a mated pair dies, the female cannot fend for the litter and herself, the litters will die and the female will disperse to find a new mate. The vacated range will soon be filled by young, dispersing black-backed jackals and the boundaries of the existing ranges are adjusted. 10. The black-backed jackal has an opportunistic, omnivorous diet. It hunts and scavenges alone or in pairs for food but occasionally forms packs to hunt larger antelopes that are old, weak, sick or injured. Carcasses are detected by an acute sense of smell. 11. The first food resource that is encountered when foraging will be utilised and the presence of abundant natural prey will therefore buffer livestock to some extent against depredation. 12. Prey is run down and killed with a bite to the side of the neck, below the eyes or on the throat. Tooth puncture marks are 21 to 30 cm apart and the kill is opened at the groin. Occasionally the carcass is pulled away from the kill site. 13. Black-backed jackals prey heavily on herbivorous springhares, hares, mice and rats whenever they are available and consequently provide more plant food for small, domesticated livestock and other herbivores. 14. In a sheep production region in KwaZulu-Natal black-backed jackals have been estimated to be responsible for the loss of 0,05% of the sheep population. 15. Depredation on the lambs of antelope and small, domesticated livestock is seasonal and happens when the lambs are small. Scats that contain the remains of small, domesticated livestock do not indicate whether the animal was freshly scavenged or hunted and may give a skewed impression. 16. The reproductive system of the black-backed jackal is one of obligatory monogamy and synchronized breeding in females. Litters of up to nine pups are born in the winter and early spring and fertility is high. Consequently a black-backed jackal population can recover rapidly from carnivore control programmes or epizootics.

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17. Young, dispersing jackals can move for distances of more than 100 km away from the natal site to occupy vacant ranges from an age of 11 months but the life expectancy in the wild is unknown. 18. Among carnivores, black-backed jackals are preyed on by eagles, caracals, spotted hyenas, leopards and lions, but are usually competitively dominant over caracals. Black-backed jackals in turn prey on a variety of smaller carnivores and the kittens of the caracal. 19. Its acute sense of smell makes the baiting and trapping of black-backed jackals difficult. The black-backed jackal is wary and cunning which often stops it from entering a trap or taking bait. 20. Once a black-backed jackal has been involved in an unsuccessful attempt to trap or bait it, it will seldom repeat the experience and further trapping or baiting for control becomes almost impossible. 21. To date all attempts at the control of black-backed jackal populations have failed. Holistic ecosystem husbandry of small, domesticated livestock is a viable alternative. Sterilisation of breeding pairs may have some chance of success but will be difficult to achieve economically and practically. 22. Lethal approaches to the control of black-backed jackals include the use of coyote getters, packs of hunting hounds, leg-hold (gin) traps and digging out pups from dens to kill them. Except the latter one, all the methods are unselective in killing the target animal and will also kill many types of non-target animal. 23. Hunting jackals from the air with the aid of helicopters is only effective for some animals that have not yet become so shy of the sound of a helicopter that they will hide or change their activity cycle following persecution. When hunting ecologically similar coyotes from a helicopter in the USA, helicopter-shy coyotes and a change in their activity cycle eventually required the use of a back-up fixed wing aircraft and a ground crew of hunters. 24. At most, hunts from a helicopter should be confined to short periods of control just before the lambing season(s). The success rate obtained will also depend on the terrain, the density of the black-backed jackals, their experience with helicopter hunts, the type of helicopter, the hunting experience and technique of the crew and the timing and season of hunting. 25. Data from 287 hunts of black-backed jackals with helicopters have shown that 62% of the stomachs contained the remains of domesticated sheep. However, the unit costs per jackal should be calculated and compared with means of preventing the predation of jackals on small, domesticated livestock.

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26. Oranjejag, one of the largest and oldest carnivore hunting groups in South Africa, could not succeed in controlling the black-backed jackal population in their region of operations over several decades of intensive effort. Their control efforts were unselective and they also killed almost three times as many Cape foxes and numerous African wildcats. In KwaZulu-Natal, the population of black-backed jackals stayed stable despite 15 consecutive years of hunting jackals with trained hounds. 26. Leg-hold (gin) traps are unspecific in trapping carnivores, cause trauma because of pain and injury and usually end in a lingering death. Their use has sparked considerable moral debate. Leg-hold traps that kill specific types of carnivore are legal but their use becomes illegal when a threatened or endangered species is killed. Soft-catch and pan tension variations are more selective, but the method remains labour-intensive and requires proper training. 27. Shooting black-backed jackals with the aid of spotlights is not an option because these jackals avoid persecution and bright lights. 28. Poisoned baits that are laced with strychnine and Compound 1080 are at times used to control black-backed jackals during rabies epizootics but they create wide-ranging ecological problems because they are unselective. Some poisons are strictly controlled substances because they are lethal to humans. Strychnine is the only poison that can currently be obtained under prescription from a pharmacy in South Africa. 29. Coyotes-getters are unselective because they use a baited barrel that is loaded with a cartridge containing sodium cyanide to attract a carnivore and will kill most types of scavenging carnivore in South Africa. Most getters are only triggered once because a specific territorial animal has been killed. However, animals that are not killed initially will not return to a getter that has discharged close to their faces. Long-term control programmes with getters are a waste of manpower, but short-term control can be effective. 30. Coyote-getters can be used for the short-term removal of habitual killers of small, domesticated livestock when they are set around carnivore-proof overnight and maternity pens. 31. Young black-backed jackals can be dug out of dens and killed but total control with this method is impossible to achieve because not all the dens are usually found. This method is therefore uneconomical and ineffective in the control of the black-backed jackal. 32. Poisoned collars can be fitted to domesticated livestock to kill any carnivore that attacks them. Such collars can be used in combination with carnivore-proof herding and overnight and maternity pens. The poison being used is lethal to humans and should be treated with care.

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33. Shock collars that are combined with baited shock stations have been tested experimentally on wolves in the USA. Their use requires a substantial investment in labour and infrastructure and the trapping of a carnivore to be able to fit the collar. Therefore its use is considered to be impractical and expensive. 34. The only effective programme to control black-backed jackal numbers would be to exterminate all the jackals nationally. To do this is economically impossible and unsound ecologically. Moreover, the role of black-backed jackals as predators of small, domesticated livestock will also in all likelihood be taken over by another type of carnivore, such as the caracal, should the jackals be exterminated. 35. Natural predator defence behaviour can be bred into small, domesticated livestock by selectively removing those ewes immediately from the flock who fail to protect their lambs against attack by a carnivore. 36. An ecosystem approach to animal husbandry that is combined with the use of livestock protection or herding guard dogs is considered to be the best option to limit the depredation of carnivores on small, domesticated livestock. The short-term use of coyote-getters or other methods to reduce carnivore populations locally just before the lambing season(s) may also be considered. 37. When using livestock protection or herding guard dogs it requires patience, maintenance and understanding to train each dog and immediate results should not be expected. 38. Common non-lethal methods to prevent depredation include re-establishing defence behaviour against carnivores in a flock through selective breeding, the use of contraceptive baits, using cage traps to capture and re-establish carnivores elsewhere, the use of aversion or repellent chemicals, warning collars that are linked to a cell phone and scaring methods such as loud noises that are transmitted intermittently. 39. Following control, a black-backed jackal population can be expected to increase again rapidly. 40. An ecosystem husbandry approach to small, domesticated livestock production will buffer flocks against depredation because it supplies an abundant alternative food source to opportunistically feeding carnivores.

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The caracal 1. The caracal is a member of the mammal family Felidae that diverged from the phylogenetic line of the other carnivores some 40 million years ago. Caracals have existed for 2,5 million years and caracal fossils show that they lived alongside early humans at Swartkrans in the Gauteng province from two to one million years ago, 1,5 million years ago at Hopefield in the Western Cape province and more recently at the Cave of Hearths near Makado in the Limpopo province. 2. The caracal is not a type of lynx but it is an ecologically similar, agile and slender cat of medium size which can jump prodigious distances. 3. The caracal is a monotypic cat species that occurs throughout Africa, the Arabian Peninsula, Middle-East and India. 4. The caracal has a catholic habitat preference but it does favour semi-arid woodlands, wetlands and grasslands with dense scrub. It occurs in commercial plantations but avoids dense forests. 5. The caracal is mainly active at night but it may become active by day in protected areas. It climbs trees well. 6. The range of an adult male includes that of several adult females whose range size varies with habitat quality and a variable density of prey. In the semi-arid Karoo the range of an adult male can be 440 km2 or more. The boundaries of the range are delimited by scent marks. 7. Young caracals disperse after becoming nine months old and soon fill ranges that become vacant because of natural mortality or caracal control. Dispersal can take place over a distance of 65 km from the natal site or more. 8. The caracal is mainly solitary, except when mating, although a female and her kittens will move around together. 9. Hunting is done alone and the caracal mainly eats fresh prey although it will cache fresh food in a tree or under grass to return to it later. It occasionally returns to scavenge the remains of small, domesticated livestock but it sometimes also scavenges the fresh kills of other carnivores. 10. The prey is killed by a throat or nape bite after stalking and a short rush. The puncture marks of the teeth are 26 to 30 mm apart. The caracal sometimes leaves a clear imprint of its claw grip marks on the back of a kill or on the shoulders of the carcass. The caracal can bite more severely than the black-backed jackal.

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11. The carcass of a kill is generally not pulled away from the kill site and feathers and woolly fur are removed by the incisors to reach the meat. Feeding starts between the buttocks around the anus, at the shoulder or the neck. The caracal does not eat the stomach, intestines, some pieces of skin and the portion of the skull containing the teeth. 12. The caracal is independent of surface water and it eats wild fruits with high moisture content. 13. Small carnivores are killed, while birds are easily plucked from the sky with jumps of up to 2 m high. Surplus killing of small, domesticated livestock may happen especially when the livestock are kept in a pen that is not carnivore proof or are trapped against a fence. Such killing is done randomly and instinctively and such carcasses are not fed on. 14. The caracal is a generalist feeder and they will kill small, domesticated livestock. However, the main food resource consists of small rodents, hyraxes, springhares and small antelope. Adult females with kittens mainly feed on rodents. The small herbivorous prey animals being killed are an ecological bonus to the producer of small, domesticated livestock as it increases the stocking capacity of the habitat for larger herbivores. 15. Hyraxes are preyed upon in a density-dependent way and the caracal is the main regulator of hyrax populations in the Karoo. An adult caracal requires some 1 kg of food per day. 16. Caracals co-exists with several other small types of carnivore but are dominated ecologically by the black-backed jackal, although the caracal does kill the pups of the black-backed jackal. 17. In the southern Kalahari the caracal mainly killed sheep during the cold season lambing season when their natural prey base was reduced but springhares were the most common prey. 18. The stomachs of caracals that are killed in small, domesticated sheep production regions show a variable evidence of sheep remains but sheep never form the major food source. 19. Creating a healthy natural food source through an ecosystem approach to the husbandry of small, domesticated livestock will buffer the impact of the caracal on such livestock. 20. The caracal reproduces throughout the year from an age of 12 to 15 months in a male and 14 to 16 months in a female. The births peak from October to February in South Africa and litters of up to six kittens are born in caves, or in the cavities in trees or the ground. These cavities are lined with feathers or fur.

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21. A female caracal can reproduce until the end of her life expectancy at 19 years of age. 22. The caracal preys upon several types of small carnivore including black-backed jackal pups and is preyed upon in turn by the lion, leopard and spotted hyena. The black-backed jackal will prey on caracal kittens and following large black-backed jackal eradication programmes the caracal population in an area will increase rapidly. 23. In 1989 it was estimated that 2219 caracals were being killed in the Karoo region per year at considerable cost in terms of manpower and money. Yet caracal control programmes are being continued 22 years later. 24. Aerial hunting of the caracal is almost impossible because cats are wary of aircraft and will hide when hearing them approach. Aerial hunting of the ecologically similar bobcat in the USA is being done by using a specially modified fixed-wing aircraft that at times may be combined with a helicopter which elevates the costs. 25. The caracal only forms 11,5% of the 287 black-backed jackals and caracal that were hunted from the air in the examined records, but 42,2% of these stomachs contained the remains of domesticated sheep. 26. Caracals are difficult to trap and the trapping success only is around 2%. This makes trapping as a control measure uneconomical. However, when they are to be trapped the urine of a female in oestrus, or the fresh prey of a caracal can be used as bait. Baits that contain fish meal and decomposed blood and meat are sometimes successful too. 27. Expandable poisoned collars that contain carbofuran or Compound 1080 and are fitted to the lambs of small, domesticated livestock will kill attacking caracals. However, Compound 1080 is a banned substance because it is also lethal to humans, while carbofuran that is spilled will kill scavengers. 28. Hunting caracals with packs of trained hunting hounds is unselective. The Oranjejag hunting club used leg-hold (gin traps) and such hounds to attempt to control caracal numbers but it was unsuccessful although they killed 3377 caracals in 32 years of operation. The peak number of caracals was killed in the final year (1991) of recorded operations. 29. Some depredation by caracals on small, domesticated livestock can be prevented by following an ecosystem-based husbandry approach that ensures the creation of a sustainable prey resource especially during the lambing season(s). Such depredation can also be prevented by using livestock protection or guard herding dogs and other guard animals such as the donkey and llama.

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30. Repellent collars that prevent caracals from attacking small, domesticated livestock may be unsuitable for caracals because a caracal can bite through them as it has a strong jaw structure and musculature. Bell collars and those that contain aversion chemicals that cause irrational fear or dislike may also work. A collar that activates a distress call on a mobile phone has also been developed and used. 31. Herding small, domesticated livestock in overnight and maternity pens that are carnivore-proof will prevent depredation by caracals. The depredation impact of caracals can also be reduced through the selective breeding of ewes that show natural defence against carnivore attacks on their lambs. 32. Calling and shooting caracals in professional and recreational shooting has had limited success in reducing depredation by the ecologically similar lynx elsewhere and that was considered to have no practical application in controlling lynx numbers. 33. The depredation by the caracal on small, domesticated livestock can only be stopped by total extermination of the caracal on a national scale at huge economic cost and is impractical. General conclusions 1. The management of carnivore depredation on small, domesticated livestock has been practised for centuries but the control of wild carnivore populations has failed in all the documented cases that did not involve the total eradication of the carnivores. 2. The cost of carnivore control measures is usually only measured in terms of the number of carnivores being killed while it should be measured relative to the true costs of the damage and control and the benefits that are being gained from alternative options. 3. Financial and ecological management plans should be developed that balance the loss of livestock though depredation against the benefits of taking an ecosystem-based approach to the husbandry of small, domesticated livestock. 4. Submission of financial and ecological management plans for small, domesticated livestock production could become the point of departure for conservation authorities in evaluating applications for carnivore control permits. 5. The black-backed jackal and caracal are the products of a long period of development and co-existence with humans and are adapted to it. It is

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impossible to control their population sizes except through regional or national extermination. 6. The fences on small, domesticated livestock production units are usually not effective in limiting the movements of the black-backed jackal and caracal. 7. It seems to be more prudent ecologically and economically to maintain functional ecosystems on small, domesticated livestock production units and to take preventative measures to reduce or exclude the contact between such livestock and carnivores. 8. Husbandry practices for the production of small, domesticated livestock cannot be ecologically and economically sustainable when they are only driven by short-term financial gains. 9. The lambs and lambing season(s) of small, domesticated livestock are critical components that can be protected by taking measures that prevent depredation. This sometimes only requires small adjustments to the husbandry approach that is being used. 10. Controlling the impact of black-backed jackals and caracals on small, domesticated livestock cannot be done merely through attempts at controlling the carnivore population sizes. A multi-faceted approach is required that is aimed at minimizing their depredation impact. 11. The minimization of depredation by the black-backed jackal caracal on small, domesticated livestock can be achieved by taking an ecosystem-based approach to their husbandry in combination with using preventative methods to limit the contact between these livestock and carnivores. 12. The ecological benefits of the black-backed jackal and caracal on functional ecosystems include an increased habitat capacity to support herbivorous animals. This is a bonus that should form part of the overall financial balance statement. 13. The uncontrolled use of leg-hold (gin) traps and poisoned bait are non-specific and hazardous to humans, livestock and biodiversity conservation. Upon the removal of some carnivores from an ecosystem their ecological role will be taken over by other carnivores. 14. The control of carnivores does not reduce their numbers permanently because of the resilience of natural populations. Mortalities that are caused by carnivore control operations will replace those that would have been caused naturally.

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15. The indiscriminate killing of carnivores on most production units of small, domesticated has not stopped the impact of the black-backed jackal and caracal after decades of control. 16. A carefully balanced approach to limit or prevent depredation on small, domesticated livestock should be followed that is based on ecosystem-based husbandry and selected preventative measures of depredation. Specific conclusions and recommendations 1. The black-backed jackal and the caracal are the products of a long period of development alongside humans. 2. The wide geographic distribution and rapid population recovery rate make it ecologically and economically impossible to eradicate the black-backed jackal and caracal locally and regionally. 3. All attempts to control black-backed jackal and caracal populations through intensive control programmes over the past several decades in South Africa have failed. 4. Lethal control mechanisms are largely unspecific and should only be used for the short-term reduction of black-backed jackal and caracal numbers during the lambing season(s). 5. Methods to prevent or limit depredation on small, domesticated livestock at critical periods are the only option that is viable over the long term. 6. The benefits and costs of an ecosystem-based approach to the husbandry of small, domesticated livestock should be included in any such programme. 7. Predation on small, domesticated livestock by habituated carnivores can be countered by increasing ecosystem functioning in combination with using carnivore-proof pens and livestock protection or herding guard dogs. 8. Lethal methods can be used to reduce carnivore populations before the lambing season(s) but their use requires professionally trained staff. 9. Short-term permits for carnivore control should only be granted to small, domesticated livestock producers who are following an ecosystem-based approach to husbandry in combination with methods to prevent depredation. Applications for such permits should be accompanied by viable ecological and financial management plans.

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10. It seems to be time for a unified approach in which all the parties that are involved will have to yield some ground so that they all may eventually benefit from developing a new and holistic strategy. In doing so, limited agendas will have to be traded for broader visions in a spirit of true cooperation. 11. Conservation authorities cannot adopt an approach that will lead to the degradation of biodiversity and is only aimed at the economic benefits of livestock production because that is not their mandate. 12. It is suggested that conservation authorities facilitate a series of research projects concerning the ecology and control of the black-backed jackal and caracal.

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LITERATURE REVIEW

The following is a general review of the principles that are involved in carnivore ecology and carnivore-prey relationships. It is followed by a more specific review of the literature on the ecology and control of the black-backed jackal Canis mesomelas and the caracal Caracal caracal in South Africa, with reference to relevant studies elsewhere in the world. Full scientific names are given when an organism is first mentioned in the text to avoid confusion between common names. 1. Carnivore ecology The primary origin of any carnivore conflict with humans is competition for the same resources because humans and their domesticated livestock occur with various carnivores in space and time. The carnivore ancestors that first moved to Africa some 30 to 25 million years ago were the civet-like ancestors of some of the cats. They were followed later by the ancestors of the bears, dogs and some other cats. The current carnivores of Africa are all 5 million years or less old and have only come into contact with pastoralists and their domesticated livestock in the past 10 000 to 12 000 years. However, they have been associated with humans for millions of years. Conflict arises because the domesticated livestock have often lost those instinctive defence mechanisms against carnivores which enabled their ancestors to avoid or accommodate predation. A further source of conflict is that carnivores are by nature hunters or scavengers that eat meat. All carnivores occupy ranges that are large enough to support them and their young. These ranges are demarcated by scent marks and the ranges are usually defended against intrusion by others of the same kind through social behaviour. On commercial livestock production units or farms which may have become denuded of such natural prey this defence behaviour may become diluted. The size of a range depends on the availability of prey animals and is smaller in wetter regions with a high incidence of prey than in arid regions where prey is scarce. Young animals disperse after becoming independent to find ranges that have been vacated as a result of natural mortalities, the latter which include predation among carnivores. Such young animals are the most highly productive element of any population and will result in rapid population growth. The only way in which the carnivore impact on any prey be minimized is by effective exclusion of depredation or by killing all the predators that move into a given region in a blanket national or provincial extermination campaign. The latter is ecologically undesirable and economically impossible to do. Moreover, numerous protected areas and those in which no control is possible will act as reservoirs for repopulation through the dispersal of young animals. There are no documented successful extermination cases involving any of the smaller carnivores, although some of the larger carnivores are currently under conservation threat following merciless persecution. These larger carnivores also

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require ranges that are too large to be maintained in most regions of the modern world. Carnivores will utilise the most abundant prey resource that is available to them and will switch to alternative prey when their main prey become scarce. Moreover, they all hunt opportunistically and will usually kill the first suitable prey animal that they come across. Maintaining a healthy population of natural prey on a small, domesticated livestock production unit will therefore act as a buffer system against depredation on the livestock. For the smaller carnivores such buffer prey are usually smaller mammals, many of which may have a vegetarian diet. Carnivores cannot regulate the population of any prey animal that is abundant, but will do so if a prey resource becomes limited. Some carnivores occasionally become habituated to target a specific prey resource; such as unprotected small, domesticated livestock; that is abundant and easy to kill. Only the guilty animal should then be removed because the random killing of various types of carnivore can do more ecological harm than good. When a specific carnivore learns to attack and kill small, domesticated livestock, such a habitual killer is usually an adult, territorial male but not all the adult territorial males in a carnivore population will become habitual killers. Control measures that are unselective place the producers of small, domesticated livestock who use them in a possible untenable legal position because of the regulations that have been issued in 2007 for the National Environmental Biodiversity Act (Act 10 of 2004) prohibit the killing of rare wildlife that may become targeted. In functional, large conservation areas the wild herbivore herds are able to maintain substantial production levels despite the presence of large carnivores because they display natural defence mechanisms against predation. Where such defence mechanisms have been lost, livestock husbandry that is based on ecosystem principles and the re-introduction of such defence mechanisms through selective breeding can yield positive results. There also is evidence that carnivores will target sick or weak animals in preference to healthy ones. Whenever depredation by a specific carnivore is suspected, the first step is to ensure that the correct cause of death has been confirmed. Livestock may die of various causes, and because most carnivores are also scavengers any carcass may attract them. The presence, position, spacing and depth of teeth puncture marks will indicate if and which carnivore may have been responsible for the death of an animal. The method of killing is also important as will be explained below for the black-backed jackal and the caracal. Other indicators include the feeding pattern, the nature and presence or not of drag marks, and bleeding patterns.

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In South Africa, the lion Panthera leo, brown hyaena Hyaena brunnea and the spotted hyaena Crocuta crocuta are the only sociable carnivores that live in large prides and clans. The other carnivores, such as the black-backed jackal and caracal, live singly or at most in pairs and will never reach high densities at a given place and time. Because of ecosystem collapse on most of the small, domesticated livestock production units, the main smaller carnivores that are responsible for most livestock losses in South Africa are the black-backed jackal that lives in mated pairs and the single-living caracal. Moreover, their predation impact is usually most severe during the autumn and spring lambing season(s) of the livestock. Carnivores can be divided broadly into two energy balance models based on the energetic requirements of their feeding strategies. Small-bodied carnivores, such as the black-backed jackal and the caracal, that weigh <20 kg, will mainly feed on prey that are smaller than themselves, while the large-bodied carnivores mainly feed on large prey. Relatively large, but still small-bodied, carnivores such as a large caracal can obtain more energy by occasionally switching to larger prey, but such a prey would require twice as much energy to hunt as a small prey-specialist of equivalent body size. The ultimate body size of a modern carnivore is that of the polar bear Ursus maritimes in which an adult weighs around 1000 kg. The smaller carnivores that weigh <14,5 kg, such as the black-backed jackal, will prefer to hunt smaller prey which are energetically more profitable. Their size and prey preferences make the larger carnivores more vulnerable to extinction pressures and the same will apply to smaller carnivores that hunt larger prey. Forcing them to do so through small, domesticated livestock production programmes which do not follow healthy ecosystem principles and are only aimed at economic gain can expose them and the livestock producers to long-term risk through ecosystem collapse. Predation is a vital part of ecosystem functioning and the dynamics of carnivore populations cannot be divorced from broader environmental issues. Fully functional ecosystems with at least a healthy carnivore complement form crucial base-lines against which to measure ecological change and productivity. This is also true for the husbandry of small, domesticated livestock on an ecosystem management basis where there is an interaction between humans, carnivores and their prey (including small, domesticated livestock). However, generalizations are invalid and every situation must be studied in a particular environment at a specific time. In some cases, carnivores that only kill specific types of prey because of skewed habitat management programmes will ultimately limit the production potential of the prey, including small, domesticated livestock. When an ecosystem management approach is being used in the husbandry of small, domesticated livestock it can be beneficial to livestock and wildlife production because of the interspecific competition between different types of

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carnivore. For example, in the Yellowstone area of the USA, the presence of increasing numbers of the gray wolf Canis lupus lupus have led to larger populations of pronghorn antelope Antilocapra americana because the wolves were killing and scaring off coyotes Canis latrans that were preying on the pronghorn antelope lambs, while the wolves were preying on larger prey than the pronghorn antelope. A cascade of effects may follow the extinction of carnivores from ecosystems, with subsequent alteration of the habitat and even destructive effects on bird populations. In the Karoo region of South Africa, the production of small, domesticated livestock over ten years has caused changes in the distribution of the small mammals and the structure of the vegetation. Following coyote control on small, domesticated livestock production units in the USA, wild ungulates may become numerous and cause overgrazing. The net result is overall lower productivity and a loss of biodiversity. A large natural ecosystem such as the Serengeti is self-sustaining through food availability and carnivore-prey interactions provided that human-induced change is kept to a minimum. The best approach therefore seems to be to manage all land on an ecosystem basis. On conservation areas this implies that carnivore management has to be based on a programme of no intervention. Many popular perceptions that exist among the public and some media on livestock-carnivore interactions are not supported by the results of a multitude of studies by independent and highly rated scientists over many decades all over South Africa and elsewhere. Yet the livestock producers and the media usually agree that livestock production methods as practised in the past have disturbed natural carnivore-prey and ecosystem balances that have developed over centuries. As a consequence these imbalances have become a contributory factor to creating the current conflict between livestock producers and carnivores. There also is an increasing move towards the use of non-lethal interventions in carnivore management. The economic benefits of following an holistic ecosystem approach to the production of small, domesticated livestock and wildlife can be enormous as is illustrated by one such livestock and wildlife producer from the Venterstad region in the Karoo who has been following this approach (see Smith 2011 below). Bibliography Anon 2008. Researchers say more gray wolves mean more pronghorn antelope in the Yellowstone area. Associated Press, 3 March. Berger, J 2007. Fear, human shields and the redistribution of prey and predators in protected areas. Biology Letters 3: 620 – 623.

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Berger, K M 2006. Carnivore-livestock conflicts: effects of subsidized predator control and economic correlates on the sheep industry. Conservation Biology 20(3): 751 – 761. Bothma, J du P. 2010. Predators: general principles. In J du P Bothma and J G du Toit (Eds), Game ranch management, fifth edition. Pretoria: Van Schaik, pp 289 – 295. Carbone, C, A Teacher and J M Rowcliffe 2007. The costs of carnivory. PLoS Biology 5(2): 363 – 368. Eccard, J A, R B Walther and S J Milton 2000. How livestock grazing affects vegetation structure and small mammal distribution in the semi-arid Karoo. Journal of Arid Environments 46: 103 – 106. Harrington, J T and M R Conover 2007. Does removing coyotes for livestock protection benefit free-ranging ungulates? Journal of Wildlife Management 71(5): 1555 – 1560. MacRae, C 1999. Life etched in stone. Johannesburg: Geological Society of South Africa. Roberts, D H 1986. Determination of predators responsible for killing livestock. South African Journal of Wildlife Research 16: 150 – 152. Sinclair, A R E, S A R Mduma, J G C Hopcraft, J M Fryxell, R Hilborn and S Thirgood 2007. Long-term ecosystem dynamics in the Serengeti: lessons for conservation. Conservation Biology 21(3): 580 – 590. Smith, M 2011. Vee en wild wei hier saam. Landbouweekblad, 8 July: 4 – 6. Snow, T V 2008. A systems-thinking based evaluation of predator conflict management on selected South African farms. Master’s degree in Environment and Development dissertation. Pietermaritzburg: University of KwaZulu-Natal. Stuart, C T 1985. Reading animal sign. The Naturalist 29: 6 – 20. Treves, A and U Karanth 2003. Human-wildlife conflict and perspectives on carnivore management worldwide. Conservation Biology 17(6): 1491 – 1499. Van Rooyen, M 2011. Jakkals-taakspan “tydmors”. Rapport, 6 February: 17.

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2. Carnivore-prey interactions and carnivore control Large carnivores are not always detrimental to other animals and predation is a vital part of ecosystem functioning. Carnivore-prey interactions in a large, natural ecosystem such as the Serengeti are self-regulating provided that human-induced changes are kept to a minimum. Therefore the dynamics of carnivore-prey interactions cannot be divorced from broader environmental principles and issues. Exploitative competition for food is wide-spread among carnivores and is ecologically and economically important. The intensive removal or absence of one type of carnivore will influence the population dynamics of another one, and hence also ecosystem dynamics. Detrimental cascading effects may follow the extinction of carnivores from ecosystems, with subsequent alteration of the habitat and even destructive effects on productivity and bird populations. The average carnivore in Africa is prone to predation and influence, whether directly as prey or through interspecific competition, by 15 other types of carnivore at some stage of its life. Carnivores in a natural system cannot regulate the numbers of abundant prey, but they do regulate the numbers of the smaller resident ungulates and the small carnivores. Carnivore-livestock predation management has been practised for centuries but except for total extermination of carnivores it has failed in all properly documented cases. Moreover, the benefits and costs of control measures are usually being measured only through the number of carnivores being killed, while a more recent systems analysis has indicated that it should be measured in proportion to the true costs of the damage or loss and the benefits gained. This will require more careful financial analyses, with the loss of livestock as a result of predation and the positive spin-off of taking an ecosystem management approach to livestock husbandry being built into financial management plans. Ecosystem management programmes should include data on trends in the carnivore populations and the productivity of the habitat to support herbivores over time. Even small adaptations towards the improved husbandry of small, domesticated livestock may make a substantial difference to the impact of their depredation and the productivity of the habitat to support herbivores. Ecological and financial management plans should therefore include information on the long-term trend in the success, costs and benefits of any carnivore control programme. Inclusion and the evaluation of an ecological and financial management programme could become the point of departure for conservation authorities when considering applications for carnivore control permits. In such applications, the combination of landscape and livestock management variables will have to be used to explain patterns of depredation and the need for carnivore control. Subsidization of the livestock industry from government funds to compensate livestock producers for losses that are caused by carnivores is sometimes muted,

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but in North America it has not produced positive results in maintaining or increasing sheep production. Bibliography Anon 2008. Researchers say more gray wolves mean more pronghorn antelope in the Yellowstone area. Associated Press, 3 March. Berger, J, P B Stacey, L Bellis and M P Johnson 2008. A mammalian predator-prey imbalance: grizzly bear and wolf extinction affect avian Neotropical migrants. Ecological Applications 11(4): 947 – 960. Berger, K M 2006. Carnivore-livestock conflicts: effects of subsidized predator control and economic correlates on the sheep industry. Conservation Biology 20(3): 751 – 761. Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. Conradie, B, J Piesse and C Thirtle 2009. What is the appropriate level of aggregation for productivity indices? Comparing district, regional and national measures. Agrekon 48(1): 9 – 20. Linnell, J D C, J Odden, M E Smith, R Aanes and J T Swenson 1999. Do “problem individuals” really exist? Wildlife Society Bulletin 27(3): 698 – 705. Michalski, F, R L Boulhosa, A Faria and C A Peres 2006. Human-wildlife conflicts in a fragmented Amazonian forest landscape: determinants of large felid depredation on livestock. Animal Conservation 9: 179 – 188. Sinclair, A R E, S A R Mduma, J G C Hopcraft, J M Fryxell, R Hilborn and S Thirgood 2007. Long-term ecosystem dynamics in the Serengeti: lessons for conservation. Conservation Biology 21(3): 580 – 590. Smith, M 2011. Vee en wild wei hier saam. Landbouweekblad, 8 July: 4 – 6. Smuts, G L 1978. Interrelations between predators, prey and their environment. BioScience 28(5): 316 – 319. Snow, T V 2008. A systems-thinking based evaluation of predator conflict management on selected South African farms. Master’s degree in Environment and Development dissertation. Pietermaritzburg: University of KwaZulu-Natal. Woodroffe, R, L G Frank, P A Lindsey, S M K ole Ranah and S Romañach 2007. Livestock husbandry as a tool for carnivore conservation in Africa’s community rangelands: a case-control study. Biodiversity Conservation 16: 1245 – 1260.

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3. The value of scientific nomenclature In dealing with and analysing depredation on small, domesticated livestock and carnivore control it is important first to ensure that the individual and the type of carnivore that is involved be identified clearly. The type of animal that is responsible for depredation can be identified based on its killing and feeding techniques, but the age and sex of animal that is responsible for the depredation should also be established so as not to target all the members of their kind in control or preventative programmes. When identifying the type of animal that is involved, it should be recognized that there are numerous regional and even localised common names for animals. For the black-backed jackal, for example, there are at least 16 different common names in Afrikaans, only one of the 11 official languages in South Africa. Moreover, the same common name is sometimes being used for different types of animal. It therefore becomes difficult and confusing to determine to which type of animal a specific reference really refers and consequently leads to misconceptions and hearsay perceptions. To deal with this confusion and the proliferation of common names, a binary system of scientific nomenclature was designed in the 1750s by Carl Linnaeus who classified all the known organisms of the world into various taxonomic categories (taxa) in a series of volumes under the title Systema Naturae. This system of scientific names should be used to verify the origin of any depredation on small, domesticated livestock. The first volume of Systema Naturae deals with the broad classification of biota into plants, animals and other life forms. The animals are broadly classified into those animals with internal skeletons (vertebrates) and those without them (invertebrates). The vertebrates are classified into five classes to represent the Amphibia (amphibians), Reptilia (reptiles), Pisces (fishes), Aves (birds) and Mammalia (mammals). The mammals are in turn classified into Orders, Suborders, Infraorders, Parvorders, Superfamilies, Families, Subfamilies, Tribes, Genera, Species and Subspecies. The scientific names of the genera, species and subspecies are indicated in italics in text and are the real level of use to identify various types of mammal. Subspecies are isolated geographically and reproductively but they can interbreed when this isolation is lost. The oldest scientific name has preference except when it is based on a misconception. For instance, the greater kudu Tragelaphus strepsiceros was originally described as Antilope strepsiceros by Pallas in 1766 but because the genus Antilope was later found only to occur as the blackbuck Antilope cervicapra in India, the generic name Tragelaphus was created in 1816 by De Blainville for the species of this genus in Africa. On the species level Linnaeus created a binary system within which organisms are first classified into broad genera, such as the dog Canis, and then as different species such as the black-backed jackal Canis mesomelas. Some genera such as Caracal and Antidorcas are monotypic because they only consist of a single

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species. Moreover, some types of animal are also subspecies as they have already diverged into geographically or reproductively isolated populations, such as the black-backed jackal Canis mesomelas mesomelas. Bibliography Bothma, J du P 1993. Probleemdiere slagoffers van naamsverskille. Landbouweekblad, 24 December: 18 – 21. Stuart, C T 1985. Reading animal sign. The Naturalist 29: 6 – 20. Wilson, D E and D M Reeder (Eds), Mammal species of the world – a taxonomic and geographic reference, third edition. Baltimore: Johns Hopkins University Press. 4. The black-backed jackal The black-backed jackal is a member of the wide-spread family Canidae that includes animals that are ecologically adapted so well that they colonized most of the Earth. There are eleven species of canid in Africa. The fossil evidence indicates that black-backed jackals have diverged from other jackals as much as 2,3 to 2,5 million years ago. Fossilized black-backed jackals are known to have lived alongside early humans some 2 million years ago in Kenya and Tanzania. In South Africa they are known from fossils at Swartkrans in the Gauteng province where they lived some 2 to 1 million years ago. The name jackal is derived from Persian name sagal which probably refers to the golden jackal Canis aureus. In Swahili the black-backed jackal is named jasira which means bold and courageous. Long before the time of the European settlers, the oral history of the Khoikhoi indicates that they had already experienced depredation by black-backed jackals on their small, domesticated livestock. The black-backed jackal was first described scientifically as Canis mesomelas by Schreber in 1775 based on a specimen that was collected in the Western Cape province of South Africa. In South Africa it is a carnivore of medium size, but the size varies by region. The weight of an adult male ranges from 5,9 to 12,0 kg and that of a female from 5,5 to 10,0 kg. The black, saddle-like markings on the back give this animal its most often used common name. Age determination can be done on the basis of the relative pulp width of the canine teeth which will distinguish young from adult animals. Bibliography Anon 2011. The black-backed jackal. http://en.wikipedia.org/wiki?Black-backedJackal

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Brain, C K 1993. Swartkrans: a cave’s chronicle of early man. Transvaal Museum Monograph 8: 1 – 270. Bingham, J and G K Purchase 2002. Age determination in jackals (Canis adustus Sundevall, 1846, and Canis mesomelas, Schreber, 1778: Carnivora: Canidae) with reference to the age structure and breeding patterns of jackal populations in Zimbabwe. African Zoology 38(1): 153 – 160. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491. Wozencraft, W C 2005. Order Carnivora. In D E Wilson and D M Reeder (Eds), Mammal species of the world – a taxonomic and geographic reference, third edition. Baltimore: Johns Hopkins University Press, pp 532 – 628. 4.1 Ecology The black-backed jackal weighs less than the side-striped jackal Canis adustus that occurs in the high rainfall parts of north-eastern South Africa and further north into Africa. The side-striped jackal is not much of a scavenger and seldom, if ever, attacks small, domesticated livestock. In any programme of reducing the impact of black-backed jackals on small domesticated livestock, the following ecological parameters have to be taken into consideration as some perceptions about black-backed jackals are based only on hearsay and has no factual basis. 4.1.1 Distribution As does the bat-eared fox Otocyon megalotis, the black-backed jackal occurs as two distinct subspecies in southern and eastern Africa, with Canis mesomelas mesomelas in southern Africa and Canis mesomelas schmidti in eastern Africa. These two subspecies are both known as black-backed jackals but they are currently separated by a wetter belt of habitat which is at least 900 km wide although the species most likely had an unbroken distribution in an earlier more arid period in the history of Africa. In South Africa the black-backed jackal is ubiquitous as it occurs in all regions, extending north into southern Africa as far as south-western Angola, most of Botswana, south-central Zimbabwe and southern Mozambique. Bibliography Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491.

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4.1.2 Habitat preference Being found throughout South Africa, the habitat of a black-backed jackal there varies widely but there is some preference for open woodlands and grasslands. Nevertheless it occurs from the semi-arid western regions to the wetter eastern ones with a rainfall of up to 1000 mm per year. It is only absent from deep forests. In the north-eastern parts of the Limpopo province of South Africa and southern Mozambique it occurs in more open habitats while the heavier side-striped jackal occurs in the river valleys. Black-backed jackals rest underground in burrows, in rock crevices or among boulders in rocky areas. Near human habitation they rest in open areas that provide them with a clear view of the surrounding terrain. Bibliography Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. Rowe-Rowe, D T 1992. The carnivores of Natal. Pietermaritzburg: Natal Parks Board. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491. 4.1.3 Range use and activity patterns As in other canids, range use by black-backed jackals is greatly influenced by the topography, vegetative cover, the availability of food in time and space and human disturbances. In functional ecosystems these ranges are much smaller than in many commercial livestock-producing units which are largely devoid of natural cover and food. Consequently the range sizes vary from 1,3 to 575 km2

depending on local circumstances. The overall mean territory size for a mated adult pair is 10,6 km2 (1060 ha) in agricultural areas, but it is 18,2 km2 in the Giant’s Castle Game Reserve in KwaZulu-Natal, 4,2 km2 in the semi-arid Kalahari ecosystem and a minimum of 24,9 km2 in the Namib Desert. Because of the extremely low food resources, territoriality does not occur in the Namib Desert where the ranges of mated adult pairs overlap to a large extent. Mated pairs will also occasionally make forays into neighbouring territories in search of food. Nevertheless, a mated pair of black-backed jackals is usually almost exclusively territorial, although neighbouring territories can overlap by up to 10%, which limits overpopulation. Territories are marked by scent marks to alert other jackals that these territories are occupied and this creates a natural spacing pattern. Territory defence breaks down temporarily at an abundant food resource such as a large carcass and a large number of black-backed jackals may gather at such a carcass. Such a

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gathering is often mistakenly interpreted as proof of a high density of jackals. Young jackals that do not become helpers disperse over distances of 5 to 135 km way from their natal sites to find vacant territories. This limits the effectiveness of localised black-backed jackal control programmes. Black-backed jackals in protected areas are active by day and night and their activity pattern closely follows that of their main murid (mice) prey. However, activity is reduced in bright sunshine and on moonless nights when vision is impaired. In areas that are more densely populated by humans, or where they are being persecuted actively, black-backed jackals are mainly active at night, sunset or sunrise. A study that was done in the USA showed that the activity pattern of the ecologically similar coyote can be influenced by the type of disturbance which the animal experiences. Bibliography Bothma, J du P 1971. Notes on movement by the black-backed jackal and the aardwolf in the western Transvaal. Zoologica Africana 6(2): 205 – 207. Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer, pp 7 – 14. Ferguson, J W H, J A J Nel and M J de Wet 1983. Social organization and movement patterns of black-backed jackals Canis mesomelas in South Africa. Journal of Zoology (London) 199: 487 – 502. Ferguson, J W H, J S Galpin and M J de Wet 1988. Factors affecting the activity patterns of black-backed jackals Canis mesomelas. Journal of Zoology (London) 214: 55 – 59. Grobler, J H, A Hall-Martin and C Walker 1984. Predators of southern Africa. Johannesburg: Macmillan, pp 20 – 21. Hayward, M W and G J Hayward 2010. Potential amplification of territorial advertisement markings by black-backed jackals (Canis mesomelas). Behaviour 147: 979 – 992. Hiscocks, K and M R Perrin 1988. Home range movements of black-backed jackals at Cape Cross Seal Reserve, Namibia. South African Journal of Wildlife Research 18(3): 97 – 100. Kaunda, S K K 2000. Activity patterns of black-backed jackals at Mokolodi Nature Reserve, Botswana. South African Journal of Wildlife Research 30(4): 157 – 162.

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Kitchen, A M, E M Gese and E Schauster 2000. Changes in coyote activity patterns due to reduced exposure to human persecution. Canadian Journal of Zoology 78: 853 – 857. Moehlman, P D 1978. Jackals. Wildlife News, Kenya 13: 1 – 6. Rowe-Rowe, D T. 1982. Home range and movements of black-backed jackals in an African montane region. South African Journal of Wildlife Research 12: 79 – 84. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 420 – 424. 4.1.4 Social structure Canids are unique among carnivorous mammals in that the members of a group will share food and care for sick adults and dependent young. The black-backed jackal, however, does not live in large groups. It mates for life with the mated pair being the centre of a social unit that defends a territory aggressively against other jackals and limits overpopulation. The boundaries of a territory are scent-marked and the black-backed jackal is highly vocal. The use of territories prevents overpopulation by jackals in any given area because the territory size is related to prey abundance. The young of the previous litter may occasionally help to care for and feed the young. When one member of a mated pair dies, the other cannot fend for itself and the litter and the female may die or the female will disperse. However, the vacant territory will be filled almost immediately by a young and vigorously reproducing pair while the boundaries of the existing ranges are adjusted. In black-backed jackals this social segregation may break down temporarily at an abundant food source such as a large carcass and all the pairs in a given region may congregate at such a food source while it is still available. For example, a large number of jackals may gather at an elephant carcass, while on the Skeleton Coast in Namibia, where black-backed jackal pairs are not strictly territorial, 78 jackals have been seen to aggregate at a seal colony. Carcasses are located by the acute sense of smell of a black-backed jackal. Bibliography Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer, pp 7 – 14.

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Ferguson, J W H 1978. Social interactions of black-backed jackals Canis mesomelas in the Kalahari Gemsbok National Park. Koedoe 21: 151 – 162. Ferguson, J W H, J A J Nel and M J de Wet 1983. Social organization and movement patterns of black-backed jackals Canis mesomelas in South Africa. Journal of Zoology (London) 199: 487 – 502. Hayward, M W and G J Hayward 2010. Potential amplification of territorial advertisement markings by black-backed jackals (Canis mesomelas). Behaviour 147: 979 – 992. McKenzie, A A 1990. Co-operative hunting in the black-backed jackal Canis mesomelas. PhD thesis. Pretoria: University of Pretoria. McKenzie, A A 1993. Biology of the black-backed jackal Canis mesomelas with reference to rabies. Onderstepoort Journal of Veterinary Research 60: 367 – 371. Nel, J A J and R Loutit 1986. The diet of black-backed jackals (Canis mesomelas) along the Namib Desert Coast. Cimbebasia Series A 8: 91 - 96. Oosthuizen, W H, M A Meÿer, J H M David, N M Summers, P G H Kotze, S W Swanson and P D Shaughnessy 1997. Variation in jackal numbers at the Van Rheenen Bay seal colony with comment on likely importance of jackals as predators. South African Journal of Wildlife Research 27: 26 – 29. Smithers, R H N 1971. The mammals of Botswana. Museum Memoirs 4. Salisbury: Trustees of the National Museums of Rhodesia, 147 – 151. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 420 – 424. 4.1.5 Diet and killing method As in all canids, black-backed jackals are coursing carnivores that run their prey down before killing them. They forage singly but in the Mashatu Game Reserve in Botswana they may form temporary packs of up to 12 jackals to hunt adult impalas Aepyceros melampus that are old, weak, sick or injured. In KwaZulu-Natal, 78% of 872 sightings were of single jackals. Prey that lack defences against predators are therefore easy to kill. The prey size is consistent with the size of a canid. As in other carnivores the diet of a black-backed jackal varies regionally based upon the food resource that is available. In conservation areas such as the Serengeti ecosystem, jackals often kill the lambs of the smaller antelope but the black-backed jackals there still scavenge 35% of their food.

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When hunting in packs a herd of antelope is rushed to separate weak or injured animals which are then run down much as a Cape wild dog Lycaon pictus pack does. All the impalas that were killed in this way in the Mashatu Game Reserve in Botswana, however, were found to be old, in an extremely poor condition or had been injured previously. Pack-hunting therefore only happens when prey have become vulnerable. Moreover, the jackals will stop hunting when the prey runs off. A prey animal is killed with a bite to the side of the neck, under the eyes or the throat. The tooth puncture marks are usually some 21 to 30 mm apart, the carcass is opened at the groin and the intestines and buttock are eaten first. Cartilage and thin bones are also eaten and the carcass is usually pulled away from the site where it was killed. Springbok Antidorcas marsupialis remains were found abundantly in 313 and 522 scats that were collected respectively on two private wildlife ranches in the Northern Cape Province. However, there was no evidence on whether the springbok were hunted or scavenged, or on whether they were young or older individuals. The black-backed jackal is an omnivorous opportunist and eats a variety of vertebrate animal and plant food, with carrion, insects and warm-blooded prey being eaten most often. Black-backed jackals are likely to use the first food resources that become available once they become active. They are attracted by the smell of the placentas of antelope and sheep when they give birth and this makes the lambs secondary targets. In protected areas, antelope lambs are common prey although a black-backed jackal can kill larger prey such as adult antelopes and will scavenge for food as much as it would hunt fresh prey. Black-backed jackals prey heavily upon springhares Pedetes capensis, hares Lepus spp and rodents in protected areas and will readily eat wild fruits and even grapes. In one study in Botswana they mainly ate grasshoppers, crickets and termites. In a sheep production area in KwaZulu-Natal, black-backed jackals were found to be the cause death of 0,05% of the entire sheep population, mostly young lambs. In another study also in a sheep producing area in KwaZulu-Natal, sheep formed 35% of the diet of the black-backed jackals, with antelope being the main prey. In a sheep-farming region the Northern Cape province, wild mammals formed 57%, vegetable matter 19,7%, sheep 14,9% and carrion 13,9% of the diet. Rodents formed 16,5%, birds 12,7% and invertebrates 7,9% of the diet. In a high-rainfall region in KwaZulu-Natal almost half of the diet consisted of murids (mice and rats). Predation on sheep lambs is generally seasonal and mainly occurs in the winter when natural food sources become scarce. In the Orange Free State an analysis of the stomach contents of black-backed jackals showed that they mainly contained rodents and small livestock remains. The latter was expected as the stomachs were collected during jackal control operations. Moreover, the predation on small, domesticated livestock mainly occurred in the late summer. A similar analysis of 425 jackal stomachs that were

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also mainly obtained through various jackal control programmes on small, domesticated livestock production units in the former Western Transvaal, with a few from elsewhere including 17% from conservation areas, showed that 11% of the stomachs were empty. Of the remaining 378 stomachs that contained food, the stomach content volume was 27 times that of the Cape fox Vulpes chama. Of the stomachs examined, 23,8% contained the remains of domesticated sheep, while the other major food items consisted of insects (30,7%) rodents (28,8%), hares (7,6%) and plant food (55%). By volume the major food items were: carrion (25,2%), sheep (25%) and rodents (10,1%). The rodents consisted mostly of rats and mice (71,9% of all rodent food by volume), springhares (10.4%) and rodent moles (4,2%). The largest stomach contents by volume were recorded in the summer months. This analysis compared well with an earlier analysis of 201 stomachs of black-backed jackals from the south-western and central parts of the former Transvaal. In the central Namib Desert an analysis of 772 scats of black-backed jackals showed that they predominantly preyed on birds along the coast and ate plant material in the interior regions. Along the coast, seal remains and beetles were most abundant in the scats that were examined. The above studies underline the opportunistic feeding pattern of black-backed jackals. Bibliography Anon 2011. The black-backed jackal. http://en.wikipedia.org/wiki?Black-backedJackal Bothma, J du P 1965. Random observations on the food habits of certain Carnivora in South Africa. Fauna and Flora 16: 16 – 22. Bothma, J du P 1971. Food of Canis mesomelas in South Africa. Zoologica Africana 6(2): 195 – 203. Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. Bothma, J du P, J A J Nel and A Macdonald 1984. Food niche separation between four sympatric Namib Desert carnivores. Journal of Zoology (London) 202: 327 – 340. Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer, pp 26 – 32. Goldenberg, M, F Goldenberg, S M Funk, J Henschel and E Millesi 2010. Diet composition of black-backed jackals, Canis mesomelas in the Namib Desert. Folia Zoologica 59(2): 93 – 101.

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Grafton, R N 1965. Food of the black-backed jackal: a preliminary report. Zoologica Africana 1(1): 41 – 53. Kaunda, S K K and J D Skinner 2003. Black-backed jackal diet at Mokolodi Nature Reserve, Botswana. African Journal of Ecology 41: 39 – 46. Klare, U, J F Kamler, U Stenkewitz and D W Macdonald 2009. Diet, prey selection, and predation impact of black-backed jackals in South Africa. Journal of Wildlife Management 74(5): 1030 – 1042. Kok, O B 1996. Dieetsamestelling van enkele karnivoorsoorte in die Vrystaat, Suid-Afrika. South African Journal of Science 92: 393 – 398. Lamprecht, J 1978. On the diet, foraging behaviour and interspecific food competition of jackals in the Serengeti. Zeitschrift für Säugetierkunde 43: 210 – 223. McKenzie, A A 1990. Co-operative hunting in the black-backed jackal Canis mesomelas. PhD thesis. Pretoria: University of Pretoria. Nel, J A J and R Loutit 1986. The diet of black-backed jackals (Canis mesomelas) along the Namib Desert Coast. Cimbebasia Series A 8: 91 – 96. Nel, J A J, R Loutit and J du P Bothma 1997. Prey use by black-backed jackals along a desert coast. South African Journal of Wildlife Research 27(3): 100 – 104. Oosthuizen, W H, M A Meyer, J H M David, N M Summers, P G H Kotze, S W Swanson and P D Shaughnessy 1997. Variation in jackal numbers at the Van Rheenen Bay seal colony with comment on likely importance of jackals as predators. South African Journal of Wildlife Research 27: 26 – 29. Roberts, D H 1986. Determination of predators responsible for killing livestock. South African Journal of Wildlife Research 16: 150 – 152. Rowe-Rowe, D T 1975. Predation by black-backed jackals in a sheep-framing region of Natal. Journal of the southern African Wildlife Management Association 5: 79 – 81. Rowe-Rowe, D T 1976. Food of the black-backed jackal in nature conservation and farming areas in Natal. East African Wildlife Journal 14: 345 – 348. Rowe-Rowe, D T. 1982. Home range and movements of black-backed jackals in an African montane region. South African Journal of Wildlife Research 12: 79 – 84.

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Rowe-Rowe, D T 1983. Black-backed jackal diet in relation to food availability in the Natal Drakensberg. South African Journal of Wildlife Research 13: 17 – 23. Rowe-Rowe, D T 1984. Black-backed jackal population structure in the Natal Drakensberg. Lammergeyer 32: 1 – 7. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 420 – 424. Stuart, C T 1976. Diet of the black-backed jackal Canis mesomelas in the central Namib Desert, South West Africa. Zoologica Africana 11(1): 193 – 205. Stuart, C T 1985. Reading animal sign. The Naturalist 29: 6 – 20. Stuart, C T 1987. A comparison of the food of the black-backed jackal and caracal. The Naturalist 31(3): 41 – 42. Wyman, J 1967. The jackals of the Serengeti. Animals (London): 10: 79 – 83. 4.1.6 Population characteristics The mating system of the black-backed jackal is long-term obligatory monogamy (mated pairs) although mate changes do happen. The overall population sex ratio favours females and this reflects the physiological and ecological constraints of the environment. Young are born in the winter and early spring but the breeding season varies with local conditions. Mating occurs from July to October with the females showing synchronised breeding. Fertility in black-backed jackals is high but sterile mating is known. The litter size can be up to nine pups per litter. The pups are born in the winter to early spring after a gestation period of 60 to 70 days. The pups are born in an underground den and parental care is critical for pup survival. Reproduction is vigorous when a population has declined for some reason and with the large litter sizes it allows rapid recovery of a population. A female cannot raise a litter on her own if the male is killed. The male and helpers feed the lactating female and the pups are totally dependent on their parents and possible helpers for their first four months of life. The helpers are the young of the previous year. In the Serengeti ecosystem, 73% of all the breeding pairs are assisted by helpers who also protect the pups against possible predation from other carnivores and which increases the pup survival rate. In turn, the helpers have a better survival rate because of their association with the parental pair. The pups first emerge from the birth den at three weeks of age and

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young jackals become sexually mature when they are 11 months old. When young black-backed jackals are a year old they either become helpers or disperse to establish mated pairs in vacant territories. The life expectancy in the wild varies regionally but the exact parameters are unknown. Bibliography Bingham, J and G K Purchase 2002. Reproduction in the jackals Canis adustus Sundevall, 1846 and Canis mesomelas Schreber, 1778 (Carnivora: Canidae), in Zimbabwe. African Zoology 37(1): 21 – 26. Bothma, J du P 1971. Control and ecology of the black-backed jackal Canis mesomelas in the Transvaal. Zoologica Africana 6(2): 187 – 189. Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. De Vos, V 1970. Pseudo pregnancy in the black-backed jackal (Canis mesomelas Schreber). Journal of the South African Veterinary and Medical Association 40(4): 381 – 383. Fairall, N 1968. The reproductive seasons of some mammals in the Kruger National Park. Zoologica Africana 3: 189 – 210. Grobler, J H, A Hall-Martin and C Walker 1984. Predators of southern Africa. Johannesburg: Macmillan. pp 20 – 21. Moehlman, P D 1979. Jackal helpers and pup survival. Nature 277: 382 – 383. Moehlman, P D 1989. Intraspecific variation in canid social systems. In J L Gittleman (Ed), Carnivore behavior, ecology and evolution. London: Chapman and Hall, pp 143 – 163. Rowe-Rowe, D T 1975. Predation by black-backed jackals in a sheep-framing region of Natal. Journal of the southern African Wildlife Management Association 5: 79 – 81. Rowe- Rowe, D T 1984. Black-backed jackal population structure in the Natal Drakensberg. Lammergeyer 32: 1 – 7. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491.

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4.1.7 Interaction with other carnivores The black-backed jackal is prey to a host of other predators, including eagles, spotted hyenas, lions and leopards Panthera pardus. In conservation areas, spotted hyenas will raid the dens of black-backed jackals to kill and eat the pups. In the Kalahari, black-backed jackals are killed and eaten by lions and especially often by leopards. The black-backed jackal is a favourite food item of leopards in many areas, while it is also preyed upon by caracals. In the Kalahari, black-backed jackals are the target of 22% of all leopard hunts and some leopards learn to eat them exclusively. The habit of a black-backed jackal to follow a large predator while barking at it often leads to it being killed and eaten. Black-backed jackals sometimes facilitate stalking by cheetahs Acinonyx jubatus in the Nairobi National Park by finding prey and then running between the prey and the cheetahs. Black-backed jackals are superior competitors to caracals and will prey on caracal kittens. The eradication of most of their natural predators has undoubtedly advanced the expansion of black-backed jackal populations on many production units for small, domesticated livestock. Bibliography Bothma, J du P 1965. Random observations on the food habits of certain carnivora in South Africa. Fauna and Flora 16: 16 – 22. Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer, 9 – 42. Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. Bothma, J du P and E A N le Riche 1984. Aspects of the ecology and the behaviour of the leopard Panthera pardus in the Kalahari Desert. Supplement to Koedoe 1984: 259 – 279. Bothma, J du P and E A N le Riche 1986. Prey preference and hunting efficiency of the Kalahari Desert leopard. In S D Miller and D D Everett (Eds), Cats of the world: biology, conservation and management. Washington, DC: National Wildlife Federation, pp 389 – 414. Bothma, J du P and E A N le Riche 1994. Scat analysis and aspects of defecation in northern Cape leopards. South African Journal of Wildlife Research 24(1&2): 21 – 25. Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik. Eloff, F C 1973. Lion predation in the Kalahari Gemsbok National Park. Journal of the southern African Wildlife Management Association 3(2): 59 – 63.

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Grobler, J H 1981. Feeding behaviour of the caracal Felis caracal in the Mountain Zebra National Park. South African Journal of Zoology 16: 259 – 262. Moehlman, P D 1989. Intraspecific variation in canid social systems. In J L Gittleman (Ed), Carnivore behavior, ecology and evolution. London: Chapman and Hall, pp 143 – 163. Müller-Using, D 1975. Jackals. In B Grzimek (Ed), Grzimek’s animal life encyclopaedia. New York: Van Nostrand Reinhold, pp 236 – 254. Palmer, R and N Fairall 1988. Caracal and African wild cat diet in the Karoo National Park. South African Journal of Wildlife Research 18(1): 30 – 34. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 420 – 424. Stuart, C T 1987. A comparison of the food of the black-backed jackal and caracal. The Naturalist 31(3): 41 – 42. Wyman, J 1967. The jackals of the Serengeti. Animals (London): 10: 79 – 83. 4.2 Control Its partially scavenging feeding habit and an acute sense of smell are the two aspects that form the basis for black-backed jackal control. Conversely it also makes baiting and capturing this animal in traps difficult because the black-backed jackal is wary and cunning. The wily nature of a black-backed jackal also means that once a specific method of capture or control has failed, such an individual is not likely to repeat the experience. It is consequently impossible to obtain total control or extermination. Nevertheless, many techniques have been developed over time but there is an increasing focus on non-lethal methods. Bibliography Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491. Treves, A and U Karanth 2003. Human-wildlife conflict and perspectives on carnivore management worldwide. Conservation Biology 17(6): 1491 – 1499.

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4.2.1 The nature and effectiveness of control measures Black-backed jackals have been subjected to sustained control measures over many decades, all of which have failed and are labour-intensive. Some require trained personnel and cannot be applied personally by livestock producers. Many lethal methods have been developed to control black-backed jackal numbers, but in all documented cases some of the jackals learned to avoid these devices, usually following some malfunction of the equipment, and hence made future control ineffective. A study in Zimbabwe has indicated that black-backed jackal populations are capable of rapid recovery following the death or removal of even a large portion of a population. Therefore, a carefully balanced approach that includes ecosystem management and the prevention of depredation has to be considered. After review of the coyote control measures in The USA it has been suggested that the indiscriminate killing of carnivores such as the coyote, and hence the ecologically similar black-backed jackal, is not a feasible approach to limit the impact of carnivores on small, domesticated livestock. Another study concluded that the sterilisation of breeding coyotes appears to offer the largest and most lasting control of coyote populations. However, whether this can ever be done economically and practically on a regional basis is doubtful. In the USA, non-lethal control includes variations in the husbandry approach to small, domesticated livestock, while most lethal control methods are unselective and aimed at the localised reduction of coyote numbers. Bibliography Blejwas, K M, B J Sacks, M M Jaeger and R McCullough 2002. The effectiveness of selective removal of breeding coyotes in reducing sheep predation. Journal of Wildlife Management 66(2): 451 – 462. Connolly, G E 1978. Predator control and coyote populations: a review of simulation models. Coyote. New York: Academic Press. Connor, M M, M R Ebinger and F F Knowlton 2008. Evaluating coyote management strategies using a spatially explicit, individual-based, socially structured population model. Ecological Modelling 219: 234 – 247. Knowlton, F F, E M Gese and M Jaeger 1999. Coyote depredation control: an interface between biology and management. Journal of Range Management 52(5): 398 – 412. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations.

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Mitchell, B R, M M Jaeger and R Barrett 2004. Coyote depredation management methods and research needs. Wildlife Society Bulletin 32(4): 1209 – 1218. 4.2.1.1 Helicopter hunts Canids are at times hunted from the air by using helicopters or fixed-wing aircraft. The use of a helicopter to shoot black-backed jackals is not a new concept and was already tested with unknown results in the Karasburg district in Namibia in 1979. The test involved combining a helicopter with 15 mounted hunters on the ground who communicated by radio with the helicopter staff. In the USA, specific benefits were obtained by hunting the coyote from the air. However, the coyote becomes wary of a helicopter and persecution over time and will then hide upon hearing the sound of a helicopter. This later necessitated that the use of helicopters had to be combined with that of fixed-wing aircraft and ground teams of hunters to remain reasonably effective. New immigrants to areas where coyotes had been controlled were expected to be more vulnerable to aerial hunting because they were unfamiliar with their new range. Changing their activity pattern when being persecuted and becoming shy of helicopters and hiding when they hear the characteristic sound of a helicopter happens in many canids and can also be expected in black-backed jackals because they are ecologically similar to coyotes. It has been found that aerial hunting of coyotes some three to six months before moving sheep into high mountain pastures in Utah and Idaho in the USA was an effective management strategy to reduce livestock losses. The hunting of coyotes from helicopters was also considered to be economical in one study in the USA when it was compared with appointing carnivore control staff on a permanent basis but the costs were not compared with the benefits of a more ecological approach to the husbandry of small, domesticated livestock. Moreover success was related to various variables such as terrain, coyote density, type of aircraft, hunting technique and the timing of aerial hunting. As this study was done some 12 years ago, it may also no longer be cost effective due to the escalating costs of operating helicopters. Black-backed jackals formed 88,5% of the 287 jackals and caracals that were killed during helicopter hunts in the records of the South West Africa Administration Division Nature Conservation and Tourism. Of these jackals, the stomachs of 62,2% contained sheep remains, 24,4% were empty, and the rest contained small numbers of hares, birds, steenbok Raphicerus campestris, springbok, mongooses, rodents and hyraxes Procavia capensis, while one contained a lizard. Two of the stomachs contained the pods of the Prosopis tree and four contained grass. Especially older black-backed jackals are likely to become helicopter shy and wise. Moreover, they switch to a nocturnal lifestyle when being persecuted.

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Flying helicopters close to the ground at night then will be prone to flying hazards, while the unit cost per kill of black-backed jackals by using helicopters, fixed-wing aircraft and hunters on the ground in combination should be compared with other methods to reduce or prevent jackal predation on small, domesticated livestock. Increasing costs and flying hazards are the main reasons why unmanned aircraft are being used increasingly as environmentally friendly legislative alternatives to helicopters and fixed-wing aircraft in wildlife management work in the USA. Bibliography Dolbeer, R A, N R Holler and D W Hawthorne 1996. Identification and control of wildlife damage. In T A Bookhout (Ed), Research and management techniques for wildlife and habitats, fifth revised edition. Bethesda: The Wildlife Society, pp 474 – 506. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Lensing, J E 1979. ‘n Eenmalige proef op die beheer van die rooijakkals (Canis mesomelas Schreber 1775) met ‘n helikopter. Windhoek: Suidwes-Afrika Administrasie Afdeling Natuurbewaring en Toerisme. Wagner, K and M R Conover 1999. Effect of preventative coyote hunting on sheep losses. Journal of Wildlife Management 63(2): 606 – 612. 4.2.1.2 Other methods Carnivore hunting clubs have been controlling black-backed jackal numbers for many years with packs of trained hunting hounds, gin traps, coyote getters and specialized staff. One of the largest such organisations was Oranjejag in the Free State. Their efforts proved to be largely ineffective at controlling the numbers of black-backed jackals partially because of the vigorous breeding of young, dispersing jackals that occupied the ranges that were left vacant when killing territorial jackals. From 1959 to 1991 the staff of Oranjejag killed 24 589 jackals (786,4 per year) but the kills varied with the quality of the packs and staff being used. The kill of 1452 jackals in the final year of records in 1991 was the best success rate ever, showing that the jackal population had not been controlled. Moreover, from 1965 to 1991 a large number of other carnivores was killed that were not responsible for small livestock depredation, including 65 415 Cape foxes (2516,1 per year) with a peak of 4329 foxes being killed in 1994. In addition, 4892 African wildcats Felis sylvestris were killed (188,2 per year) with a peak of 671 African wildcats being killed in 1968. Over the period of 32 years 3377 caracals were also killed, while dogs, both domestic and feral, formed 2945 of the kills from 1971 to 1991.

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Of all the jackals that were killed with hunting hounds from 1962 to 1966 in KwaZulu-Natal, few young jackals were ever killed, and few adults were killed from May to September when the pups were being raised. Further analysis showed that the population of black-backed jackals there had stayed stable for 15 consecutive years of hunting. It has also been shown that black-backed jackal populations recovered rapidly after being controlled during a rabies-induced population collapse. The black-backed jackal is extremely difficult to trap and although it may be attracted to a trap site by suitable bait, it will usually not enter the trap but will circle it warily without entering it. Leg-hold (gin) traps target a variety of animals in a non-specific way. Their use also evokes a considerable moral debate among the public and trained wildlife professionals because the use of such traps may cause serious injuries to animals that free themselves, while those that remain trapped die a lingering and painful death unless the traps are serviced several times per day. Shooting black-backed jackals with spotlights at night is not an option because they keep well clear of and move away from bright lights. Nevertheless they may penetrate urban areas freely and will hunt close to farm buildings. Until the 1990s, meat bait that was laced with strychnine was used to control black-backed jackal numbers during rabies outbreaks. Strychnine was classified as a Schedule 2 substance until early 1981 which meant that it could be purchased in many places without prescription. In 1981 it was rescheduled to Schedule 4 because of its malicious use in killing dogs. Hence it became a subscription substance that could only be obtained from a veterinarian or medical doctor and is now only available from a pharmacy. It also became illegal to use strychnine without a permit that was issued by the relevant provincial authority, although it remains the only legal poison to use in carnivore control. Compound 1080 (sodium mono-fluoro acetate) has also been used in baited carcasses but some scavengers have not been seriously affected by it because they avoid the poisoned parts. In its use to control black-backed jackals, the non-selective nature of poisoned bait became a serious ecological problem. Moreover, the doses that are required for different types of animal vary greatly. For example, when taken orally with bait, humans can be killed with the same dose that is required to kill a black-backed jackal. The Wildlife Group of the SA veterinary Association does not condone the use of bait that is laced with poison because the method is inhumane and also kills of a host on non-target free-living wildlife that also eat meat. Moreover, it is lethal to humans. Bait that is laced with Compound 1080 is still used in large parts of Australia to control alien red foxes Vulpes vulpes and such poisoned baits are being set out in many wilderness areas. The advantage in Australia is that there are relatively few wild carnivores that can be harmed in the process. Even when using an acceptable alternative

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poison it will require careful control and record keeping as happens with those drugs that are being used for wildlife capture. In South Africa, it was reported that in one area 113 Cape vultures Gyps coprotheres were killed from 1 March 1983 to 28 February 1984 after eating baits that contained strychnine and were intended for black-backed jackals. Other wild carnivorous animals that are known to have been killed included eagles with serious biodiversity conservation consequences. Moreover, the occurrence of rabies is not a regular phenomenon and is the product of ecological circumstances. Strychnine also has serious health side-effects on small, domesticated livestock that are poisoned by accident or design. Another serious problem is that strychnine is not easily biodegradable as it has a half-life of 50 years and should only be used during extreme rabies epizootics. The coyote-getter is a deadly control measure that uses a sodium cyanide cartridge in a short, baited barrel that is hammered into the ground. The barrel is enveloped in a thin wool cover which is coated with smelly bait that usually consists of putrid meat, fat and brains. When an animal tries to pick up the bait the trigger mechanism is activated and the cyanide is discharged into its mouth provided that the target animal had completely covered the bait with its mouth. This again is an unselective method while jackals that have had a cyanide cartridge go off close to the face when the mouth did not cover the barrel would never take this bait again, reducing its future effectiveness as a control measure. In a study in the former Transvaal province, most of the 1237 coyote-getters that were used to kill 564 jackals in one control programme were triggered within the first two weeks of setting. Moreover, 78,4% of the getters were only triggered once indicating that long control programmes with coyote-getters are a waste of manpower. A specific jackal that has become habituated to killing small, domesticated livestock was usually removed by a strategically placed getter within the first week of setting it. Hunting black-backed jackals with spotlights at night at bait as it is sometimes being done is costly and inefficient. It is also unselective as any black-backed jackal that is called up will be killed and not necessarily a jackal that has been involved in predation of small, domesticated livestock. Another possible method of control is to sterilise whole populations of jackals. A study on such sterilisation in a coyote population in the USA revealed that it reduced, but did not eliminate, coyote depredation on sheep. However, the degree of reduction obtained indicated that this could prove to be beneficial as a small-scale method of black-backed jackal control on production units of small, domesticated livestock. The use of leg-hold (gin) traps, sometimes laced with strychnine or another poison, which are placed out strategically is more effective than most other methods. However, it is also inhumane and non-selective, especially when baits are used to lure carnivorous animals indiscriminately to the trap site. In terms of the National Environmental Management Biodiversity Act (Act 10 of 2004) and its

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regulations as published in 2007 the use of gin traps is legal for the black-backed jackal. The problem lies in the unselective nature of the method as it becomes a legal transgression as soon as a threatened species is killed. The use of Soft Catch leg-hold traps and correct pan tensions can make these traps more selective, but the use of leg-hold traps involves considerable labour as the traps must be inspected several times per day to prevent injury or a lingering death. Another method targets jackals in dens by locating them, digging them out and killing them. In the ecologically similar coyote in the USA, young and non-breeding coyotes were mainly removed by this method. In black-backed jackals, this method proved to be uneconomical and ineffective as all the young jackals in a specific region could never be located and removed. Moreover, jackal pups normally remain in a den for their first three weeks of life only. The use of poisoned collars on livestock is a more selective approach as it only targets those black-backed jackals that have become habituated to killing sheep. It is also the technique that has satisfied most of the producers of small, domesticated livestock who suffer from coyote depredation on such livestock in the USA. It can be used in conjunction with overnight and maternity pens. The poison collar kills an attacking carnivore because the original collar, known as the McBride collar, contains the toxin carbofuran, which was later replaced by Compound 1080 (sodium mono-fluoro acetate), in a pouch that is punctured when a carnivore attacks a small, domesticated animal. One problem is that when it is spilled, carbofuran is extremely toxic to scavengers. Moreover, in South Africa Compound 1080 is a banned substance because among others it is also lethal to humans although it is still being used in Australia. Shock collars were used experimentally in conjunction with bait stations in the control of wolves in the USA and could have a possible application to black-backed jackals. The principle is one of behavioural conditioning to divert wolves from specific sites or areas, such as the pastures of small, domesticated livestock. Innotek Training Collars (Invisible Fence Technologies, Garrett, Indiana, USA) were used and probes make contact with the skin of the throat. Bait stations were used with a shock tower being placed in the middle and the wolves were fitted with shock collars. When the collars were activated the wolves received a low-impulse shock to the throat every 13 seconds upon entry into the shock zone. The experiment indicated that the shock collars gave a degree of non-lethal control over the movements of free-ranging wolves where lethal means of control have become ineffective. However, the shock habituation was not permanent. Based on their distribution over the entire South Africa and the long distances over which young animals disperse to occupy vacant areas, the only effective control programme would be to exterminate black-backed jackals nationally. This would be undesirable ecologically and impossible economically. The removal of carnivores could create the possibility of an explosion in natural prey, many of

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them competing with domesticated livestock for food. Moreover, upon the removal of black-backed jackals from the environment their ecological role may well be taken over by other predators, especially the caracal. Even local, and if possible regional, extermination is not an option. An ecosystem husbandry approach that is being combined with measures to prevent or control the impact and contact of black-backed jackals with small, domesticated livestock is the only viable alternative. Moreover, one sheep producer in the Venterstad area of the Karoo is also re-establishing natural defence mechanisms among his sheep by removing and selling any ewe that loses her lamb to a carnivore. Bibliography Anon 1986. Proceedings of a colloquium on the use/abuse of strychnine. Onderstepoort: The Wildlife Group of the South African Veterinary Association, 15 February. Anon 1992. Oranjejag carnivore control statistics. Orange Free State: Oranjejag. Anon 2011. The black-backed jackal. http://en.wikipedia.org/wiki?Black-backedJackal Bothma, J du P 1971. Control and ecology of the black-backed jackal Canis mesomelas in the Transvaal. Zoologica Africana 6(2): 187 – 193. Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. Bothma, J du P 2010. Aerial counts of wild animals. In J du P Bothma and J G du Toit (Eds), Game ranch management, fifth edition. Pretoria: Van Schaik, pp 467 – 476. Bromley, C and E M Gese 2001. Surgical sterilization as a method of reducing coyote predation on domestic sheep. Journal of Wildlife Management 65(3): 510 – 519. Burns, R J and G E Connolly 1995. Assessment of potential toxicity of Compound 1080 from livestock protection collars to canines and scavenging birds. International Biodeterioration and Biodegradation 1995: 161 – 167. Dolbeer, R A, N R Holler and D W Hawthorne 1996. Identification and control of wildlife damage. In T A Bookhout (Ed), Research and management techniques for wildlife and habitats, fifth revised edition. Bethesda: The Wildlife Society, pp 474 – 506. Dyer, J 1989. Bekendstelling van kleinvee-beskermingshalsband (livestock protection collar). Worcester: The Livestock Protection Co.

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Hawley, J E, T M Gehring, R N Schulz, S T Rossler and A P Wydeven 2009. Assessment of shock collars as nonlethal management for wolves in Wisconsin. Journal of Wildlife Management 73(4): 518 – 525. Kamler, J F, N F Jacobsen and D W Macdonald 2008. Efficiency and safety of Soft Catch traps for capturing black-backed jackals and excluding non-target species. South African Journal of Wildlife Research 38(2): 113 – 116. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. McKenzie, A A 1993. Biology of the black-backed jackal Canis mesomelas with reference to rabies. Onderstepoort Journal of Veterinary Research 60: 367 – 371. Mendelssohn, H. 1989. Felids in Israel. Cat News 10: 2 – 4. Muth, R M, R R Zwick, M E Mather, J F Organ, J J Daigle and S A Jonker 2006. Unnecessary source of pain and suffering or necessary management tool: attitudes of conservation professionals toward outlawing leghold traps. Wildlife Society Bulletin 34(3):706 – 715. Pringle, J A and V L Pringle 1979. Observations on the lynx Felis caracal in the Bedford district. South African Journal of Zoology 14: 1 – 4. Sacks, B N, K Blejwas and M M Jaeger 1999. Relative vulnerability of coyotes to removal methods on a Northern California ranch. Journal of Wildlife Management 63(3): 939 – 949. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 486 – 491. Smith, M 2011. Vee en wild wei hier saam. Landbouweekblad, 8 Julie: 4 – 6. Snow, T V 2008. A systems-thinking based evaluation of predator conflict management on selected South African farms. Master’s degree in Environment and Development dissertation. Pietermaritzburg: University of KwaZulu-Natal. Stuart, C T and V J Wilson 1988. The cats of southern Africa. Zimbabwe: Chipingali Wildlife Trust. Watts, A C, J H Perry, S E Smith, M A Burgess, B E Wilkinson, Z Szantoi, P G Ifju and H F Percival 2009. Small unmanned aircraft systems for low-altitude aerial surveys. Journal of Wildlife Management 74(7): 1614 – 1619.

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4.2.2 Methods to prevent depredation Non-lethal methods to prevent depredation on small, domesticated livestock include cage traps and the release of carnivores elsewhere, using ecosystem management in livestock husbandry, seasonal lambing in carnivore-proof maternity pens, the use of repellent collars and various scaring methods. The timing and duration of control measures are important. Localized, short-term control can be effective during the lambing season of domesticated and wild herbivores. In one study on the Rietvlei Nature Reserve near Pretoria such short-term control of black-backed jackals a few weeks before the blesbok Damaliscus pygargus phillipsi lambing season improved the survival rate of those blesbok lambs up to three weeks old from 37 to 85%. In the Free State, depredation by black-backed jackals on small, domesticated livestock reaches a peak in the late summer and extra care should then be taken to reduce the predation impact then. Repellent or King collars have been developed by the King brothers of the Eastern Cape province of South Africa and can be fitted to entire flocks of small livestock. Moreover, they are expandable to allow for the growth of individual animals. The collars basically make it impossible for a jackal to bite the throat of small livestock and therefore prevent killing even when there is contact. Bell, scent and taste aversion collars can also be used to prevent depredation by causing neophobia (irrational fear or dislike) in carnivores. The Veldwagter collar activates a sensor when an animal is being chased which then activates a mobile phone warning system. In a black-backed jackal population, oestrous females may possibly be prevented from reproducing without creating vacant ranges through the use of baits that contain contraceptive steroid hormones including estradiol benzoate. Although not yet tested for black-backed jackals it has, however, only proved to be partially effective in the ecologically similar coyote. Herding livestock by day and night is an option that works well. The use of human herders can be replaced or supplemented by using shepherd herding guard or livestock protection dogs that were first used in Europe some 2000 years ago. In the Pyrénées Range between France and Spain the more common breeds being used as such guard dogs are the Great Pyrénées, the Komondor and the Akbash, while Anatolian and Maremma breeds are also being used. The dogs are habituated from an age of six to eight weeks to grow up with a flock of small, domesticated livestock and will protect the livestock. They can be used in an overnight or maternity pen but the best results were obtained in the Pyrénées with open range flocks. The presence of the dogs and their alert and aggressive defence of the livestock prevent depredation losses.

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Anatolian shepherd herding guard dogs are being used increasingly for herding small, domesticated livestock in South Africa but their training requires patience and understanding. Nevertheless, in the Limpopo province their placement with flocks of small, domesticated livestock has lead to the virtual elimination of losses. Similar results have been obtained in Namibia, the USA, Norway and Finland. In Namibia, the use of these dogs is widely accepted by livestock producers. In other parts of the world, donkeys Equus asinus and llamas Lama glama are also used to guard livestock against carnivore attacks. The most serious problem is that some livestock producers expect them to be an immediate and sole solution to a depredation problem. The use of herding guard dogs could be combined with other techniques such as using overnight and maternity pens, removing carnivores that habitually kill small livestock, reducing the carnivore population just before the lambing season and providing an abundance of natural food through an ecosystem approach to livestock husbandry. Bibliography Andelt, W F 1992. Effectiveness of livestock guarding dogs for reducing predation on domestic sheep. Wildlife Society Bulletin 20(1): 55 – 62. Anon 1986. Proceedings of a colloquium on the use/abuse of strychnine. Onderstepoort: The Wildlife Group of the South African Veterinary Association, 15 February. Anon 2010. Anatolian shepherd livestock guarding dog project. Progress Report: Cheetah Outreach, December 2010. Bothma, J du P 2010. Jackals. Game & Hunt 16(2): 6 – 9. Carlson, D A and E M Gese 2009. Integrity of mating behaviours and seasonal reproduction in coyotes (Canis latrans) following treatment with estradiol benzoate. Animal Reproduction Science 117: 322 – 330. Dolbeer, R A, N R Holler and D W Hawthorne 1996. Identification and control of wildlife damage. In T A Bookhout (Ed), Research and management techniques for wildlife and habitats, fifth revised edition. Bethesda: The Wildlife Society, pp 474 – 506. Du Plessis, S S 1972. Ecology of the blesbok with special reference to productivity. Wildlife Monograph 30: 1 – 70. Gehring, T M, K C verCauteren and J Landry 2010. Livestock protection dogs in the 21st Century: is an ancient tool relevant to modern conservation challenges? BioScience 60(4): 299 – 308.

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Green, J S, R A Woodruff and R Horman 1984. Livestock guarding dogs and predator control. Rangelands 6(2): 73 – 76. Gusset, M, M J Swarner, L Mponwane, K Ketetile and J W Mc Nutt 2009. Human-wildlife conflict in northern Botswana: livestock predation by endangered African wild dog Lycaon pictus and other carnivores. Oryx 43(1): 67 – 72. Hansen, I, T Staaland and A Ringsø 2002. Patrolling with livestock guard dogs: a potential method to reduce predation on sheep. Acta Agriculturae Scandanavica, Section A, Animal Science 52: 43 – 48. Kok, O B 1996. Dieetsamestelling van enkele karnivoorsoorte in die Vrystaat, Suid-Afrika. South African Journal of Science 92: 393 – 398. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Lötter, P 2006. The use of cell phone technology to prevent stock losses: the “Veldwagter Veediefstal Alarm”. In B Daly, H Davies-Mostert, H Evans, S Friedmann, Y King, T Snow and H Stadler (Eds), Prevention is the cure. Proceedings of a workshop on holistic management of human-wildlife conflict in the agricultural sector of South Africa. Johannesburg: Endangered wildlife Trust. Marker, L M, A J Dickman and D W Macdonald. 2005. Perceived effectiveness of livestock guarding dogs placed on Namibian farms. Ecological Management 58: 329 – 336. O’Donnell, S, J K Webb and R Shine 2010. Conditioned taste aversion enhances the survival of an endangered predator imperilled by toxic invader. Journal of Applied Ecology 47: 558 – 565. Otstavel T, K A Vuori, D E Sims, A Vlaros, O Vainio and H Saloniemi 2009. The first experience of livestock guarding dogs preventing large carnivore damages in Finland. Estonian Journal of Ecology 58(3): 216 – 224. Pringle, J A and V L Pringle 1979. Observations on the lynx Felis caracal in the Bedford district. South African Journal of Zoology 14: 1 – 4. Smith, M E, J D C Linnell, J Oden and J E Swenson 2000. Review of methods to reduce livestock depredation I. Guardian animals. Acta Agriculturae Scandanavica, Section A, Animal Science 50: 279 – 290. Smith, M E, J D C Linnell, J Odden and J E Swenson 2000. Review of methods to reduce livestock depredation II. Aversive conditioning, deterrents and

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repellents. Acta Agriculturae Scandanavica, Section A, Animal Science 50: 304 – 315. Snow, T V 2008. A systems-thinking based evaluation of predator conflict management on selected South African farms. Master’s degree in Environment and Development dissertation. Pietermaritzburg: University of KwaZulu-Natal. 4.2.3 Methods to reduce the depredation impact From various studies, it is clear that concentrated control programmes for a limited time immediately preceding the lambing season of small, domesticated livestock will reduce the impact of depredation significantly as it is the period when these livestock are most at risk. When a specific black-backed jackal becomes habituated to kill sheep in pens at night or when jackals target the lambs during the lambing season the impact can be reduced by herding the sheep at night in carnivore-proof pens. Particularly stubborn jackals can be removed by using a coyote-getter around the periphery of a herding or maternity pen until the culprit has been killed, or poison collars that will only kill carnivores that attack small, domesticated livestock can be used. The use of shepherd herding guard dogs could substantially reduce the impact of black-backed jackals on flocks of small livestock even if it does not eradicate depredation completely. A study in Zimbabwe has indicated that black-backed jackal populations are capable of rapid recovery following the death or removal of a large portion of a population. The selective sterilization of breeding coyotes in the USA is also believed to be preferable to non-selective measures to reduce the impact of coyotes on sheep losses. However, it is doubtful that such an approach will be economically and practically viable on a regional basis. Bibliography Bingham, J and G K Purchase 2002. Reproduction in the jackals Canis adustus Sundevall, 1846 and Canis mesomelas Schreber, 1778 (Carnivora: Canidae), in Zimbabwe. African Zoology 37(1): 21 – 26. Blejwas, K M, B J Sacks, M M Jaeger and R McCullough 2002. The effectiveness of selective removal of breeding coyotes in reducing sheep predation. Journal of Wildlife Management 66(2): 451 – 462. Dolbeer, R A, N R Holler and D W Hawthorne 1996. Identification and control of wildlife damage. In T A Bookhout (Ed), Research and management techniques for wildlife and habitats, fifth revised edition. Bethesda: The Wildlife Society, pp 474 – 506. Du Plessis, S S 1972. Ecology of the blesbok with special reference to productivity. Wildlife Monograph 30: 1 – 70.

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Hansen, I, T Staaland and A Ringsø 2002. Patrolling with livestock guard dogs: a potential method to reduce predation on sheep. Acta Agriculturae Scandanavica, Section A, Animal Science 52: 43 – 48. Kok, O B 1996. Dieetsamestelling van enkele karnivoorsoorte in die Vrystaat, Suid-Afrika. South African Journal of Science 92: 393 – 398. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Marker, L M, A J Dickman and D W Macdonald. 2005. Perceived effectiveness of livestock guarding dogs placed on Namibian farms. Ecological Management 58: 329 – 336. 5. The caracal The family Felidae, of which the caracal is a species, diverged from the common carnivore phylogenetic line some 40 million years ago and the first species of the genus Felis existed since the early Pleistocene some 2.5 million years ago at Omo in East Africa. Based on fossils, the caracal has lived alongside early humans for a long time as fossil caracals are known from Swartkrans in the Gauteng province where they lived 2 to 1 million years ago and Hopefield in the Western Cape province where they lived some 1.5 million years ago, while younger fossils were found at the Cave of Hearths near Makado (formerly Potgietersrust) in the Limpopo province. The name caracal is thought to be derived from its Turkish name garah-gulak or its Uzbek name karakulak, both these names meaning black ear. The Russian name is indeed karakal. In Niger the caracal mainly hunts Dorcas gazelles Gazella dorcas and has therefore been named ngam ouidenanga or the gazelle cat. The caracal is often erroneously considered to be a type of lynx Lynx spp and has even once been called the caracal lynx although the lynx and the caracal are ecologically similar. However, the caracal is most closely related to the African golden cat Profelis aurata which only occurs in the rain forests of central and western Africa. As was the black-backed jackal, the caracal was first described scientifically in 1776 by Schreber who called it Felis caracal based on a specimen from near Table Mountain in the Western Cape province. The genus Caracal was created by Gray in 1843 when it was discovered that the caracal was not a member of the genus Felis. Consequently, both the black-backed jackal and the caracal were first described scientifically by the same person, and both the original type

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specimens came from the Western Cape province in South Africa. Despite its wide distribution the caracal is a monotypic species. The caracal is a slender cat of medium size with a weight that varies from 8 to 16 kg and a shoulder height of that varies from 405 to 535 cm in different regions. Its well-built hindquarters allow it to jump prodigious distances. The black-tufted ears are conspicuous against the tawny-brown to brick-red coat. Bibliography Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Brain, C K 1993. Swartkrans: a cave’s chronicle of early man. Transvaal Museum Monograph 8: 1 – 270. Nowell, K and P Jackson 1996. Wild cats. Cambridge: IUCN Publication Services. Savage, R J G 1978. Carnivora. In J Maglio and H B S Cooke (Eds) 1978. Evolution of African mammals. Cambridge: Harvard University Press, pp 249 – 267. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 397 – 401. Smithers, R H N 1971. The mammals of Botswana. Museum Memoir 4. Salisbury: Trustees of the National Museums of Rhodesia, pp 119 – 122. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria pp 381 – 385. Sunquist, M and F Sunquist 2002. Wild cats of the world. Chicago: Chicago University Press, pp 37 – 47. Turner, A 1997. The big cats and their fossil relatives: an illustrated guide to their evolution and natural history. New York: Columbia University Press. Wozencraft, W C 2005. Order Carnivora. In D E Wilson and D M Reeder (Eds), Mammal species of the world – a taxonomic and geographic reference, third edition. Baltimore: Johns Hopkins University Press, pp 532 – 628. 5.1 Ecology

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The caracal is found throughout South Africa and widely outside Africa. The following ecological parameters are relevant to its control 5.1.1 Distribution The caracal has a wide distribution and occurs in Africa, the Arabian Peninsula, the Middle East and India. In Africa it is found in all habitats except true deserts and tropical forests and inhabits habitats up to 3300 m above sea level. in South Africa it is only absent in die extreme north-western corner of the Northern Cape province and it does occur in the evergreen forests of the mountains of the Western Cape province. 5.1.2 Habitat preference The caracal has a catholic habitat choice but avoids dense forests and true deserts. It prefers semi-arid, open woodlands and grasslands but especially those with dense scrubs. The caracal tolerates arid conditions well and is often associated with wetlands. Caracals also occur in commercial forest plantations in KwaZulu-Natal. Bibliography Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Rowe-Rowe, D T 1992. The carnivores of Natal. Pietermaritzburg: Natal Parks Board. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 397 – 401. Smithers, R H N 1971. The mammals of Botswana. Museum Memoir 4. Salisbury: Trustees of the National Museums of Rhodesia, pp 119 – 122. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 381 – 385. 5.1.3 Range use and activity patterns The caracal is predominantly active nocturnally but in protected areas it can become partially active in daytime. It climbs well and is perfectly at home in trees which explains its occurrence in the temperate forests of South Africa. When an intruder approaches a caracal, it will climb to a safe place and camouflage itself remarkably well in rocks, trees or even among tufts of grass.

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The range use is not well known but it conforms to the typical felid pattern in which a male has a territory that includes the ranges of several females whose ranges overlap to varying degrees. The caracal does not cover its faeces and with spray-urination they may serve as scent marks to demarcate the range boundaries. Near Robertson in the Western Cape province the mean range size of female caracal was 18,2 km2 and 65 km2 in a male. In the Karoo the ranges are larger at around 48 km2 for a male. In the West Coast National Park the ranges of males had a mean size of 26,9 km2 which was much larger than that of the females at 7,39 km2. In the prey-poor southern part of the Kgalagadi Transfrontier National Park the range of a single male was 308,4 km2 in extent but it used a core area of 93,2 km2 for a period of 11 months. Habitat quality determines the size of the range as in al larger carnivores. A caracal study was done in the Mountain Zebra National Park (then 6535 ha large) in the 1980s to evaluate the role of this park as a breeding ground for caracals that preyed upon small, domesticated livestock on the surrounding agricultural land. The adult males had a range size of around 15,2 km2 and the adult females one of 5,5 km2 while females on the livestock units around the park had a mean range of 19.1 km2 which indicates a more prey-poor environment. The adult male caracals in the park were a mean distance of 4,0 km apart at a given time, and the females 2,2 km. In the Mountain Zebra National Park adult caracal males preferred to move around along the brush of the upper slopes, while the females with kittens preferred the dense riverine thickets with abundant rodents which formed their primary food. Adult females without kittens used the same upper slopes as the males. When a range became vacant through the death of a caracal, a young dispersing animal settled there and the range boundaries were adjusted. Such young caracals have a higher production potential than older established ones. Based on their range size and social behaviour, the Mountain Zebra National Park, as it was at that time, could only support 14 adult and 12 subadult caracals per year. Consequently a maximum of 12 subadult caracals would be available to disperse onto the adjacent farmland. Yet, the surrounding livestock producers killed 185 caracals on their land per year which indicated that the caracals were largely breeding on the agricultural land. Two young males in the Stellenbosch area of the Western Cape travelled widely and were probably dispersing to find permanent ranges, moving 18 and 65 kilometres since being radio-collared respectively. In the Mountain Zebra National Park, two young caracal males dispersed for distances of 10 and 22 km when they were seven to nine months old. Four adult male caracals in an Acacia woodland habitat on farmland in a part of Namibia that receives a mean of 472 mm of rain per year had a mean range size

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of 312,6 km2, but the individual range sizes varied greatly between different areas, with a largest range size of 439,8 km2 and a smallest one of 79,3 km2. As these range sizes were similar to what was found in the southern Kalahari, but much larger than what was found in the Mountain Zebra and West Coast National Parks in South Africa, it is clear that habitat quality determine range size in a caracal, and hence it population density. Bibliography Avenant, N L and J A J Nel 1998. Home-range use, activity, and density of caracal in relation to prey density. African Journal of Ecology 36: 347 – 359. Bothma, J du P and E A N le Riche 1994. Range use by an adult male caracal in the southern Kalahari. Koedoe 37: 105 – 108. Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer. Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Dragesco-Joffé, A 1993. La vie sauvage au Sahara. Paris: Delachaux et Niestlé. Marker, L and A Dickman 2005. Notes on the spatial ecology of caracals (Felis caracal), with particular reference to Namibian farmlands. African Journal of Ecology 43: 73 – 76. Mellen, J D 1993. A comparative analysis of scent-marking and reproductive behavior in 20 species of small cats (Felis). American Zoologist 33: 151 – 166. Melville, H I A S 2004. The behavioural ecology of the caracal in the Kgalagadi Transfrontier Park and its impact on adjacent small livestock producing units. MSc dissertation. Pretoria: University of Pretoria. Moolman, L C. 1986. Aspekte van die ekologie en gedrag van die rooikat Felis caracal Schreber, 1776 in die Bergkwagga Nasionale Park en op die omliggende plase. MSc dissertation. Pretoria: University of Pretoria. Norton, P M and A B Lawson 1985. Radio tracking of leopards and caracals in the Stellenbosch area, Cape Province. South African Journal of Wildlife Research 15: 17 – 24. Nowell, K and P Jackson 1996. Wild cats. Cambridge: IUCN Publication Services.

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Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 397 – 401. Smithers, R H N 1971. The mammals of Botswana. Museum Memoir 4. Salisbury: Trustees of the National Museums of Rhodesia, pp 119 – 122. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 381 – 385. Stuart, C T 1983. Aspects of the biology of the caracal (Felis caracal Schreber 1776) in the Cape Province, South Africa. MSc thesis. Pietermaritzburg: University of Natal. Sunquist, M and F Sunquist 2002. Wild cats of the world. Chicago: Chicago University Press pp 37 – 47. 5.1.4 Social structure The caracal is mainly a solitary animal although two adults may travel together at times. Usually, an adult male and female will only associate when mating. In a study in the Mountain Zebra National Park, 72% of the sightings were of single caracals, 19% were two adults and the rest were of females with kittens. Caracals have a varied of vocal repertoire. Bibliography Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Grobler, J H 1981. Feeding behaviour of the caracal Felis caracal in the Mountain Zebra National Park. South African Journal of Zoology 16: 259 – 262. Moolman, L C 1986. Aspekte van die ekologie en gedrag van die rooikat Felis caracal Schreber, 1776 in die Bergkwagga Nasionale Park en op omliggende plase. MSc dissertation. Pretoria: University of Pretoria. Norton, P M and A B Lawson 1985. Radio tracking of leopards and caracals in the Stellenbosch area, Cape Province. South African Journal of Wildlife Research 15: 17 – 24. Pringle, J A and V L Pringle 1979. Observations on the lynx Felis caracal in the Bedford district. South African Journal of Zoology 14: 1 – 4.

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Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 397 – 401. Smithers, R H N 1971. The mammals of Botswana. Museum Memoir 4. Salisbury: Trustees of the National Museums of Rhodesia, pp 119 – 122. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 381 – 385. Sunquist, M and F Sunquist 2002. Wild cats of the world. Chicago: Chicago University Press pp 37 – 47. 5.1.5 Diet and killing method A caracal hunts alone, mainly eats fresh prey and will return to a fresh carcass that it has killed itself. It is known to scavenge the fresh kills of other carnivores but it does so rarely. When hunting, caracals stalk to as close to the prey as possible and then launch a short attack. The prey is usually killed by a throat or nape bite but the caracal leaves clear claw grip marks on the back of the prey. The tooth puncture marks are 26 to 30 mm apart while the large diastema (gap between the incisor and molar teeth) allows the canine teeth to penetrate prey for almost their full length. Consequently, a caracal can deliver deep and lethal bites. The fur of woolly prey is plucked out with the incisors before starting to eat the meat, and feeding usually starts between the buttocks where the carcass is opened around the anus, on the shoulder and the neck. The stomach, intestines, pieces of skin and the portion of the skull containing the teeth are not eaten. On sheep, the red fur of a caracal is usually left on the wool. The carcass is not pulled away from the kill site but it may be covered with grass for later feeding bouts. Occasionally, but rarely, the prey is cached in a tree. The caracal is independent of surface water and it occasionally eats wild fruits that have high moisture content. Caracals will readily hunt other small carnivores such as the African wildcat, the Cape grey mongoose Galerella pulverulenta, the slender mongoose Garerella sanguinea, the yellow mongoose Cynictis penicillata, genets of the genus Genetta, the otter Atilax paludinosus, the polecat Ictonyx striatus and the suricate Suricata suricatta. They also have an exceptional ability to catch birds by leaping as high as 2 m into the air to knock them down with their front paws. The feathers of larger birds are plucked first before feeding starts, but smaller birds may be eaten with their feathers. Caracals at times indulge in surplus killing when small, domesticated livestock are kept in pens that are not carnivore-proof or are trapped against a fence line which prevents them from escaping. Because such carcasses are not fed on, they represent random and instinctive kills. Surplus killing is also known in lions

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and leopards. On production units of small, domesticated livestock, surplus killing is in part considered to be coupled to the lack of natural anti-predatory instincts in the domesticated animals. Adult males are most responsible for surplus killing of sheep in the Eurasian lynx Lynx lynx in Norway. While a large male caracal can and will kill small, domesticated livestock, its main food source consists of small rodents, hyraxes, springhares and smaller antelopes. In the Mountain Zebra National Park, a study in 1986 indicated that hyraxes (dassies) formed the bulk of the diet (53 % of all prey), with the mountain reedbuck Redunca folvorufula, rodents, red rock rabbits Pronolagus rupestris and scrub hares Lepus saxatilis making up the rest. Adult females with kittens mainly preyed on rodents along the riverine thickets. There was no indication of small, domesticated livestock in the diet of the caracals in the park. In the adjacent small livestock- producing areas hyraxes were the major food source (30%), followed by rodents (24%) while small, domesticated livestock formed 23% and birds 18% of the prey. The prey base of a caracal is consequently influenced by the abundance of the prey and much of the prey being taken has an ecologically beneficial impact for the producers of small, domesticated livestock because they are mainly herbivorous. Caracals in the Strandveld ecosystem of the West Coast National Park (then some 30 000 ha large) co-existed with the water mongoose Atilax paludinosus, grey mongoose and the yellow mongoose and rodents were the most common type of prey. The feeding habits are correlated with prey availability, but over time caracals are generalist feeders. When rodent populations decline from autumn to early spring the diet switches to insects, scorpions and insectivores. The caracal shows a wide dietary range and also feeds on small antelopes, hyraxes hares and other carnivores. Of the rodents being eaten in the West Coast National Park, the bush Karoo rat Otomys unisulcatus was eaten especially, while rodent moles (family Bathyergidae) and hyraxes were mainly hunted in the cold season when the smaller rodents declined in number. In the Cape Fold Mountains of the Western Cape province, hyraxes, small antelope and vlei rats Otomys irroratus are eaten mainly. In one study in the Karoo National Park (then 2900 ha large), the grey rhebok Pelea capreolus was the most common (23%) prey of the caracal, followed by the hyrax (dassie) (22%). The hyrax was used in a density-dependent way as a food resource and the caracal is considered to be the main regulator of the hyrax and rodent populations in the Karoo. Two other types of carnivore, the suricate and the polecat were also preyed upon. It is known from other studies that the caracal preys upon other carnivores such as the African wildcat, the black-backed jackal, the slender mongoose and various species of the genet. It was estimated that a single caracal killed 15 to 16 hyraxes per year in the Karoo National Park. There was no indication that the caracals from this park preyed on small, domesticated livestock. It has been estimated that an adult caracal requires 1 kg of food per day and that the caracal population in the Karoo National Park would eat 10 000

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kg of food annually. This would include 420 hyraxes or 30% of the expected annual population increase. The impact is density-dependent and would increase as the hyrax population increased. In the southern Kalahari, a study of 116 caracal scats that was augmented by spoor tracking to find kills revealed that the vast majority of the prey of caracals was small mammals. The caracal utilised 16 of 37 possible prey resources and rodents formed 60,9% of the diet. Springhares were an abundant food resource and formed 31,4% of the rodents being killed. Other carnivores formed 10,7% of the kills and domesticated sheep 6,9%, the latter only being found in the cold season in scats as far as 23,3 km from the nearest small, and domesticated livestock units in the adjacent Namibia. These sheep lambed in the cold season. Plant material was found in 20,5% of the scats that were examined, but the tsamma melon Citrullus lanatus was probably eaten for its moisture content and not as a food item. Tracking in the Kalahari revealed a hunting success rate of 10,1% and six types of carnivore being killed, including the black-backed jackal, Cape fox, bat-eared fox, African wildcat, striped polecat and yellow mongoose. It was also found that the caracal optimized its hunting efforts during the hot season to locate and raid the colonies of Brant’s whistling rats Parotomys brantsii. On the adjacent small, domesticated livestock units in Namibia 7% of the caracal scats which were examined contained the remains of Dorper sheep. The depredation on small livestock was confined to the dry season winter months. The caracals also preyed upon black-backed jackals, Cape foxes and bat-eared foxes and consequently influenced their abundance and ecological impact on small, domesticated livestock production. The stomachs of some half of the caracals that were hunted selectively with dogs in the Bedford district of the Eastern Cape province contained the remains of small, domesticated livestock, while many contained hyraxes. This sample was biased towards known or perceived livestock- killing caracals and was biased. A later larger sample showed a similar bias because it studied specifically targeted caracals. Hyraxes there also become the main natural prey resource when other natural prey is eliminated. In the Free State one study reported that caracals were mainly killing springhares (76% of stomachs examined) and mountain reedbuck (82%). In another study, domesticated sheep remains were found in 28% of all the stomachs that were examined in the Free State and it was concluded that the caracal was the main predation agent of small, domesticated livestock there. However, most predation occurred during the summer. In a study in Botswana gerbils and mice were the main food resource, but hares, springhares, partridges, lizards and impalas were also killed at times. In Israel, caracals mainly feed on hares and partridges, while mole-rats, hedgehogs

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Erinaceus concolor, Egyptian mongooses Herpestes ichneumon and vegetation were also eaten. In Asia in Turkmenistan, rodents and hares form the main diet. Creating an abundant natural food source through a healthy ecosystem approach to the production of small, domesticated livestock will buffer the impact of the caracal on livestock depredation, and can improve stocking densities for such livestock. The removal of some prey animals by a caracal is also beneficial to the livestock producer. For example, a single springhare consumes 9,891 kg of plant material on a dry mass basis per year which on one study area of 2200 ha in the North West province converts to 4,372 metric tonnes of plant material per year for estimated population of 442 springhares. On a wet mass basis these springhares therefore removed 16,130 metric tonnes of plant material per year. In the bushveld of the Nylsvley area of the Limpopo province colonies of gerbils, which form a major food source for caracals in some regions, reduced the available herbaceous plant biomass by 47% and the plant height by 34% near their colonies. Bibliography Avenant, N L 1993. The caracal, Felis caracal Schreber, 1776, as predator in the West Coast National Park. MSc dissertation. Stellenbosch: University of Stellenbosch. Avenant, N L and J A J Nel 1997. Prey use by four syntopic carnivores in a Strandveld ecosystem. South African Journal of Wildlife Research 27(3): 86 – 93. Avenant, N L and J A J Nel 2002. Among habitat variation in prey availability and use by Felis caracal. Mammalian Biology 67: 18 – 33. Bester, J L 1982. Die gedragsekologie en bestuur van die silwervos Vulpes chama (A Smith) met spesiale verwysing na die Oranje Vrystaat. MSc dissertation. Bloemfontein: University of the Orange Free State. Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer. Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Brand, D J 1989. The control of caracal (Felis caracal) and baboons (Papio ursinus) in the Cape Province with the help of mechanical means. MSc dissertation. Stellenbosch: University of Stellenbosch. Grobler, J H 1981. Feeding behaviour of the caracal Felis caracal in the Mountain Zebra National Park. South African Journal of Zoology 16: 259 – 262.

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Kok, O B 1996. Dieetsamestelling van enkele karnivoorsoorte in die Vrystaat, Suid-Afrika. South African Journal of Science 92: 393 – 398. Linnell, J D C, J Odden, M E Smith, R Aanes and J T Swenson 1999. Do “problem individuals” really exist? Wildlife Society Bulletin 27(3): 698 – 705. Lloyd, PA and J C G Millar 1981. A questionnaire survey of some of the larger mammals of the Cape Province. Bontebok 1: 1 – 58. Melville, H I A S 2004. The behavioural ecology of the caracal in the Kgalagadi Transfrontier Park and its impact on adjacent small livestock producing units. MSc dissertation. Pretoria: University of Pretoria. Melville, H I A S and J du P Bothma 2004. Prey selection by caracal in the Kgalagadi Transfrontier Park. South African Journal of Wildlife Research 34(1): 67 – 75. Melville, H I A S and J du P Bothma 2006. Using spoor counts to analyse the effect of small stock farming in Namibia on caracal density in the neighbouring Kgalagadi Transfrontier Park. Journal of Arid Environments 64: 436 – 447. Melville, H I A S and J du P Bothma 2006. Possible optimal foraging for Brant’s whistling rats by caracals in the Kgalagadi Transfrontier Park. African Zoology 41(1): 134 – 136. Moolman, L C 1984. ‘n Vergelyking van die voedingsgewoontes van die rooikat Felis caracal binne en buite die Bergkwagga Nasionale Park. Koedoe 27: 121 – 129. Moolman, L C 1986. Aspekte van die ekologie en gedrag van die rooikat Felis caracal Schreber, 1776 in die Bergkwagga Nasionale Park en op omliggende plase. MSc dissertation. Pretoria: University of Pretoria. Norton, P M, A B Lawson, A B Henley and G Avery 1986. Prey of leopards in four mountainous areas of the south-western Cape Province. South African Journal of Wildlife Research 16(2): 47 – 52. Nowell, K and P Jackson 1996. Wild cats. Cambridge: IUCN Publication Services. Odden, J, J C D Linnell, and P F Moa, I Herfindal, T Kvam and R Andersen 2002. Lynx depredation on domestic sheep in Norway. Journal of Wildlife Management 66(1): 98 – 105. Palmer, R and N Fairall 1988. Caracal and African wild cat diet in the Karoo National Park. South African Journal of Wildlife Research 18(1): 30 – 34.

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Pringle, J A and V L Pringle 1979. Observations on the lynx Felis caracal in the Bedford district. South African Journal of Zoology 14: 1 – 4. Roberts, D H 1986. Determination of predators responsible for killing livestock. South African Journal of Wildlife Research 16: 150 – 152. Shortridge, G C 1934. The mammals of South West Africa. London: Heinemann. Skinner, J D 1979. Feeding behaviour in caracal, Felis caracal. Journal of Zoology, London 189: 523 – 525. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 397 – 401. Smithers, R H N 1971. The mammals of Botswana. Museum Memoir 4. Salisbury: Trustees of the National Museums of Rhodesia, pp 119 – 122. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 381 – 385. Stuart, C T 1982. Aspects of the biology of the caracal (Felis caracal Schreber, 1776) in the Cape Province, South Africa. MSc dissertation. Pietermaritzburg: University of Natal. Stuart, C T 1983. Aspects of the biology of the caracal (Felis caracal Schreber 1776) in the Cape Province, South Africa. MSc thesis. Pietermaritzburg: University of Natal. Stuart, C T 1985. Reading animal sign. The Naturalist 29: 6 – 20. Stuart, C T 1986. The incidence of surplus killing by Panthera pardus and Felis caracal in Cape Province, South Africa. Mammalia 50: 556 – 558. Stuart, C T 1987. A comparison of the food of the black-backed jackal and caracal. The Naturalist 31(3): 41 – 42. Stuart, C T and G C Hickman 1991. Prey of caracal Felis caracal in two areas of Cape Province, South Africa. Journal of African Zoology 105: 373 – 381. Sunquist, M and F Sunquist 2002. Wild cats of the world. Chicago: Chicago University Press pp 37 – 47. Weisbein, Y and H Mendelssohn 1990. The biology and ecology of the caracal Felis caracal in the northern Aravah Valley of Israel. Cat News 12: 20 – 22.

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5.1.6 Population characteristics The caracal probably reproduces year-round from an age of around 12 to 15 months in males and 14 to 16 months in females but there is a peak in births in the from October to February in South Africa. Copulation may occur repeatedly over a period of three days and usually lasts a mean of 3,8 minutes at a time but it can be as much as 8 minutes. Caracals produce litters that vary from one to six after a mean gestation period of 79 (range: 68 to 81) days. The kittens are born in caves, tree cavities or the burrows of other animals that are lined with fur or feathers, the eyes open at an age of nine to ten days and the coat is light yellowish to reddish-brown in colour, with black markings on the face. The kittens begin to leave the den when they are about one month old, are weaned when they are three months old when they start making their own kills and they establish their own ranges when they are nine to ten months old. Females are known to be reproductively actively up to an age of 18 years and the life expectancy is 19 years. Bibliography Bernard, R T F and C T Stuart 1987. Reproduction of the caracal Felis caracal from the Cape Province of South Africa. South African Journal of Zoology 22: 177 – 182. Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Grobler, J H 1982. Growth of a male caracal kitten Felis caracal in the Mountain Zebra National Park. Koedoe 25: 117 – 119. Mellen, J D 1993. A comparative analysis of scent-marking and reproductive behavior in 20 species of small cats (Felis). American Zoologist 33: 151 – 166. Nowell, K and P Jackson 1996. Wild cats. Cambridge: IUCN Publication Services. Peters, G. 1983. Beobachtungen zum Lautgebungsverhalten des Karakal, Caracal caracal (Schreber, 1776) (Mammalia, Carnivora, Felidae). Bonn Zoologisches Beitragen 34: 107 – 127. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 397 – 401.

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Smithers, R H N 1971. The mammals of Botswana. Museum Memoir 4. Salisbury: Trustees of the National Museums of Rhodesia, pp 119 – 122. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 381 – 385. Stuart, C T and T D Stuart 1985. Age determination and development of foetal and juvenile Felis caracal Schreber, 1776. Säugetierkundliche Mitteilungen 32:217 – 229. Stuart, C T and V J Wilson 1988. The cats of southern Africa. Zimbabwe: Chipingali Wildlife Trust. Sunquist, M and F Sunquist 2002. Wild cats of the world. Chicago: Chicago University Press pp 37 – 47. Weisbein, Y and H Mendelssohn 1990. The biology and ecology of the caracal Felis caracal in the northern Aravah Valley of Israel. Cat News 12: 20 – 22. 5.1.7 Interaction with other carnivores Caracals will readily hunt other small carnivores such as the African wildcat, the Cape grey mongoose, the slender mongoose, the yellow mongoose, genets, the otter, the polecat and the suricate. Tracking caracals in the southern Kalahari revealed that the black-backed jackal, Cape fox, bat-eared fox, African wildcat, striped polecat and yellow mongoose, were being killed by caracals. In protected areas, the caracal shares its habitat with and occasionally is preyed upon by lions, leopards and spotted hyenas. It also shares its habitat with the black-backed jackal but the jackal is the superior competitor and preys upon caracal kittens. Following large-scale black-backed jackal eradication programmes in South Africa and Israel, the caracal population again increased rapidly. Bibliography Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer. Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Brand, D J 1989. The control of caracal (Felis caracal) and baboons (Papio ursinus) in the Cape Province with the help of mechanical means. MSc dissertation. Stellenbosch: University of Stellenbosch. Kok, O B 1996. Die dieetsamestelling van enkele karnivoorsoorte in die Vrystaat, Suid-Afrika. South African Journal of Science 92: 393 – 398.

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Mendelssohn, H. 1989. Felids in Israel. Cat News 10: 2 – 4. Melville, H I A S and J du P Bothma 2004. Prey selection by caracal in the Kgalagadi Transfrontier Park. South African Journal of Wildlife Research 34(1): 67 – 75. Palmer, R and N Fairall 1988. Caracal and African wild cat diet in the Karoo National Park. South African Journal of Wildlife Research 18(1): 30 – 34. Pringle, J A and V L Pringle 1979. Observations on the lynx Felis caracal in the Bedford district. South African Journal of Zoology 14: 1 – 4. Skinner, J D and C T Chimimba (Eds) 2005. The mammals of the southern African subregion, third edition. Cambridge: Cambridge University Press, pp 397 – 401. Smithers, R H N 1971. The mammals of Botswana. Museum Memoir 4. Salisbury: Trustees of the National Museums of Rhodesia, pp 119 – 122. Smithers, R H N 1983. The mammals of the southern African subregion. Pretoria: University of Pretoria, pp 381 – 385. Stuart, C T 1987. A comparison of the food of the black-backed jackal and caracal. The Naturalist 31(3): 41 – 42. Stuart, C T and V J Wilson 1988. The cats of southern Africa. Zimbabwe: Chipingali Wildlife Trust. Sunquist, M and F Sunquist 2002. Wild cats of the world. Chicago: Chicago University Press pp 37 – 47. 5.2 Control In 1989 it was estimated that a mean of 2219 caracals were being killed per year in control operations in the former Karoo at a rate of 0,02 to 1,6 caracals per 10 km2 at considerable cost in terms of finances and manpower. Yet the depredation on livestock has continued and is still being attempted 22 years later. There is an increasing move towards non-lethal methods of carnivore control in the world. Bibliography Brand, D J 1989. The control of caracal (Felis caracal) and baboons (Papio ursinus) in the Cape Province with the help of mechanical means. MSc dissertation. Stellenbosch: University of Stellenbosch.

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Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Treves, A and U Karanth 2003. Human-wildlife conflict and perspectives on carnivore management worldwide. Conservation Biology 17(6): 1491 – 1499. 5.2.1 The nature and effectiveness of control measures There are few methods to control caracals as they are not scavengers and are difficult to trap. The only attractant that has a small measure of success is the use of the urine of a female that is in oestrus or the use of fresh bait that had been killed by the caracal itself because it is known that caracals may return to continue to feed on the carcass of their own fresh kill. However, on small, domesticated livestock production units where caracals are being persecuted, they will not come back to their own kills. Bibliography Avenant, N L 1993. The caracal, Felis caracal Schreber, 1776, as predator in the West Coast National Park. MSc dissertation. Stellenbosch: University of Stellenbosch. Moolman, L C. 1986. Aspekte van die ekologie en gedrag van die rooikat Felis caracal Schreber, 1776 in die Bergkwagga Nasionale Park en op die omliggende plase. MSc dissertation. Pretoria: University of Pretoria. Stuart, C T and G C Hickman 1991. Prey of caracal Felis caracal in two areas of Cape Province, South Africa. Journal of African Zoology 105: 373 – 381. 5.2.1.1 Hunting from aircraft Hunting caracals from helicopters is difficult because cats are especially wary of aircraft and will hide when they hear one which is also the reason why cats cannot be counted from the air. In the USA, hunting cats such as the bobcat Lynx rufus is being done mainly from a specially modified, slow-speed, fixed-wing aircraft such as a Piper Cub that may at times be combined with a helicopter and ground crews. The caracal only formed 11.5% of the 287 jackals and caracals that were killed during helicopter hunts in the records of the Western Cape Nature Conservation Board. Of these few caracals the stomachs of 42,4% contained the remains of domesticated sheep, 12,1% were empty, 15,2% contained birds, 9,1% contained hyraxes and one each of the rest contained a hare, a slender mongoose, a polecat, a southern African hedgehog Atelerix frontalis and a steenbok, while two contained grass.

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Bibliography Dolbeer, R A, N R Holler and D W Hawthorne 1996. Identification and control of wildlife damage. In T A Bookhout (Ed), Research and management techniques for wildlife and habitats, fifth revised edition. Bethesda: The Wildlife Society, pp 474 – 506. 5.2.1.2 Other methods Caracals are difficult to trap and trapping success is usually low. However, in the Mountain Zebra National Park caracals that were returning to feed have been lured to box traps that were erected near a fresh, half-eaten mountain reedbuck carcass that was killed by the caracal itself. Attempted bait consisted of equal measures of fish meal and decomposed blood and ground meat, while some measure of success was achieved with trapping caracals in the south-western Kalahari with a mixture of decomposed meat, eggs and fish or the urine of a female in oestrus. However, in the south-western Kalahari the capture success was a low 1,4%. In a study in the West Coast National Park, 12 different types of bait were used, with a mixture of fish flower, decomposed brain tissue and decomposed blood being most successful as it also was for a study in the Mountain Zebra National Park. However, the trapping success remained low at 2,15% in the West Coast National Park and 1,0% in the Mountain Zebra National Park respectively. These trapping success rates are not cost effective. The low trapping success in this ecological group of smaller cats is one reason why a remote-controlled tele-injection system has been developed to capture carnivores recently. The use of poisoned collars on the lambs of small, domesticated livestock is a more selective approach as it only targets those caracals that have become habituated to killing sheep. The original collar, known as the McBride collar, contained the toxin carbofuran which was later replaced by Compound 1080 (sodium mono-fluoro acetate). When spilled, carbofuran is extremely toxic to scavengers. Moreover, in South Africa Compound 1080 is a banned substance because among others it is also lethal to humans. Caracals that were known to have killed small, domesticated livestock were once hunted with dogs in the Bedford district of the Eastern Cape province but the method was found to be unselective. Carnivore hunting clubs have been trying to control caracal numbers for many years with packs of trained hunting hounds, leg-hold (gin) traps, and specially trained field staff. According to the records of Oranjejag, the largest such club in the Free State, it proved to be largely ineffective at controlling the numbers of caracals as only 3377 caracals (105,3 per year) were killed with all these methods over a period of 32 years. Taking the costs involved, this was not a positive economic return on the investment. The kill of 233 caracals in the final year of records in 1991 was the best success per year

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that was obtained ever, showing that the caracal population had not been controlled over several decades of intensive control. Moreover, from 1965 to 1991 these packs of hounds also killed a large number of carnivores that were not responsible for small livestock depredation. Bibliography Anon 1992. Oranjejag carnivore control statistics. Orange Free State: Oranjejag. Avenant, N L 1993. The caracal, Felis caracal Schreber, 1776, as predator in the West Coast National Park. MSc dissertation. Stellenbosch: University of Stellenbosch. Bothma, J du P 2010. Aerial counts of wild animals. In J du P Bothma and J G du Toit (Eds), Game ranch management, fifth edition. Pretoria: Van Schaik, pp 467 – 476. Brand, D J 1989. The control of caracal (Felis caracal) and baboons (Papio ursinus) in the Cape Province with the help of mechanical means. MSc dissertation. Stellenbosch: University of Stellenbosch. Dolbeer, R A, N R Holler and D W Hawthorne 1996. Identification and control of wildlife damage. In T A Bookhout (Ed), Research and management techniques for wildlife and habitats, fifth revised edition. Bethesda: The Wildlife Society, pp 474 – 506. Dyer, J 1989.Bekendstelling van kleinvee-beskermingshalsband (livestock protection collar). Worcester: The Livestock Protection Co. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Melville, H I A S 2004. The behavioural ecology of the caracal in the Kgalagadi Transfrontier Park and its impact on adjacent small livestock producing units. MSc dissertation. Pretoria: University of Pretoria. Moolman, L C 1984. ‘n Vergelyking van die voedingsgewoontes van die rooikat Felis caracal binne en buite die Bergkwagga Nasionale Park. Koedoe 27: 121 – 129. Nowell, K and P Jackson 1996. Wild cats. Cambridge: IUCN Publication Services. Pringle, J A and V L Pringle 1979. Observations on the lynx Felis caracal in the Bedford district. South African Journal of Zoology 14: 1 – 4.

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Ryser, A, M Scholl, M Zwahlen, M Oetliker, M Ryser-Degiorges and U Breitenmoser 2005. A remoter-controlled teleinjection system for the low-stress capture of large mammals. Wildlife Society Bulletin 33(2): 721 – 730. Schiess-Meier, M, S Ramsauer, T Gabanapelo and B König. 2007. Livestock predation – insights from problem animal control registers in Botswana. Journal of Wildlife Management 71(4): 1267 – 1274. Snow, T V 2008. A systems-thinking based evaluation of predator conflict management on selected South African farms. Master’s degree in Environment and Development dissertation. Pietermaritzburg: University of KwaZulu-Natal. 5.2.2 Methods to prevent depredation In a study in the south-western Kalahari it was found that caracals from the Kgalagadi Transfrontier Park avoided the adjacent small, domesticated livestock farms in Namibia during the hot season when their own prey resource was abundant, and only entered these livestock units during the cold season lambing time when their own natural prey base became depleted. Consequently, depredation was only a factor for a portion of the year when additional care such as herding can be used to prevent depredation by caracals. In the Free State depredation by caracals in small livestock showed a peak during the summer lambing season. The repellent King collar that was referred to for the prevention of depredation on small, domesticated livestock by the black-backed jackal above may be less suitable for the caracal because it has a jaw structure and musculature that may allow it to bite through a King collar. Bell and scent collars can also used be also prevent depredation by causing neophobia (irrational fear or dislike) in carnivores. The Veldwagter collar activates a sensor when an animal is being chased which then activates a mobile phone warning system. The use of shepherd guarding or livestock protection dogs dates back to Europe for 2000 years and the use of such dogs to guard small livestock flocks in the Limpopo province has lead to the virtual elimination of losses. Similar results have been obtained with the protection of sheep flocks in the USA, Norway and Finland, and these dogs are widely accepted by livestock producers in Namibia. In Europe, while donkeys and llamas are also being used successfully to guard small livestock flocks against attacks by the lynx. In herding dogs the success rate varies based on the training and quality of the dogs being used, but such dogs are regarded as a viable management tool. The use of overnight and maternity pens that are caracal-proof will improve the survival of young animals as has been suggested in Botswana for leopard depredation on livestock.

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Bibliography Andelt, W F 1992. Effectiveness of livestock guarding dogs for reducing predation on domestic sheep. Wildlife Society Bulletin 20(1): 55 – 62. Anon 2010. Anatolian shepherd livestock guarding dog project. Progress Report: Cheetah Outreach, December 2010. Friedmann, Y King, T Snow and H Stadler (Eds), Prevention is the cure. Proceedings of a workshop on holistic management of human-wildlife conflict in the agricultural sector of South Africa. Johannesburg: Endangered Wildlife Trust. Gehring, T M, K C verCauteren and J Landry 2010. Livestock protection dogs in the 21st Century: is an ancient tool relevant to modern conservation challenges? BioScience 60(4): 299 – 308. Gusset, M, M J Swarner, L Mponwane, K Ketetile and J W Mc Nutt 2009. Human-wildlife conflict in northern Botswana: livestock predation by endangered African wild dog Lycaon pictus and other carnivores. Oryx 43(1): 67 – 72. Hansen, I, T Staaland and A Ringsø 2002. Patrolling with livestock guard dogs: a potential method to reduce predation on sheep. livestock depredation. Acta Agriculturae Scandanavica, Section A, Animal Science 52: 43 – 48. Herfindal, I, J D C Linnell, P F Moa, J Odden, L B Austmo and R Andersen 2005. Does recreational hunting reduce losses on domestic sheep? Journal of Wildlife Management 69(3): 1034 – 1042. Kok, O B 1996. Dieetsamestelling van enkele karnivoorsoorte in die Vrystaat, Suid-Afrika. South African Journal of Science 92: 393 – 398. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Lötter, P 2006. The use of cell phone technology to prevent stock losses: the “Veldwagter Veediefstal Alarm”. In B Daly, H Davies-Mostert, H Evans, S Friedmann, Y King, T Snow and H Stadler (Eds), Prevention is the cure. Proceedings of a workshop on holistic management of human-wildlife conflict in the agricultural sector of South Africa. Johannesburg: Endangered Wildlife Trust. Marker, L M, A J Dickman and D W Macdonald. 2005. Perceived effectiveness of livestock guarding dogs placed on Namibian farms. Ecological Management 58: 329 – 336.

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Melville, H I A S and J du P Bothma 2006. Using spoor counts to analyse the effect of small stock farming in Namibia on caracal density in the neighbouring Kgalagadi Transfrontier Park. Journal of Arid Environments 64:436 – 447. Otstavel, T, K A Vuori, D E Sims, A Vlaros, O Vainio and H Saloniemi 2009. The first experience of livestock guarding dogs preventing large carnivore damages in Finland. Estonian Journal of Ecology 58(3): 216 – 224. Smith, M E, J D C Linnell, J Odden and J E Swenson 2000. Review of methods to reduce livestock depredation. I. Guardian animals. Acta Agriculturae Scandanavica, Section A, Animal Science 50: 279 – 290. Smith, M E, J D C Linn ell, J Odden and J E Swenson 2000. Review of methods to reduce livestock depredation. II. Aversive conditioning, deterrents and repellents. Acta Agriculturae Scandanavica, Section A, Animal Science 50: 304 – 315. Snow, T V 2008. A systems-thinking based evaluation of predator conflict management on selected South African farms. Master’s degree in Environment and Development dissertation. Pietermaritzburg: University of KwaZulu-Natal. 5.2.3 Methods to reduce the depredation impact A study that was done in the Karoo National Park indicated the positive economic value of the caracal in controlling small herbivorous mammals and black-backed jackals to some degree. This principle could be applied on small, domesticated livestock production areas by taking an ecosystem approach to animal husbandry. It is argued that most carnivores will kill livestock at some time in their lives and that the only way in which to prevent such depredation is to remove as many individuals as possible. However, in reality, as for black-backed jackals, caracal depredation on small, domesticated livestock can only be prevented by the extermination of the caracal on a national scale with huge economic and ecological costs. Calling caracal with commercial calls and then shooting them professionally or as recreational shooting is an option that has been tried for similar-sized cats that were responsible for livestock losses in the USA and Norway. In the latter study, however, the recreational shooting of the Eurasian lynx only had a small beneficial impact on preventing sheep losses and was considered to have no practical benefit. Bibliography Hansen, I, T Staaland and A Ringsø 2002. Patrolling with livestock guard dogs: a potential method to reduce predation on sheep. Acta Agriculturae Scandanavica, Section A, Animal Science 52: 43 – 48.

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Herfindal, I, J D C Linnell, P F Moa, J Odden, L B Austmo and R Andersen 2005. Does recreational hunting reduce losses on domestic sheep? Journal of Wildlife Management 69(3): 1034 – 1042. Lamarque, F, J Anderson, R Ferguson, K Lagrange, Y Osei-Owusu and L Bakker 2009. Human-wildlife conflict in Africa. Rome: Food and Agriculture Organization of the United Nations. Linnell, J D C, J Odden, M E Smith, R Aanes and J T Swenson 1999. Do “problem individuals” really exist? Wildlife Society Bulletin 27(3): 698 – 705. Marker, L M, A J Dickman and D W Macdonald. 2005. Perceived effectiveness of livestock guarding dogs placed on Namibian farms. Ecological Management 58: 329 – 336 . Palmer, R and N Fairall 1988. Caracal and African wild cat diet in the Karoo National Park. South African Journal of Wildlife Research 18(1): 30 – 34. 6. General conclusions Carnivore-livestock depredation management has been practised for centuries, but the control of wild carnivore populations has failed in all properly documented cases. The cost of control measures is usually measured in terms of the number of carnivores being killed, while a more recent systems analysis has indicated that it should be measured relative to the true costs of the damage and control and the benefits that are being gained from such operations and alternative options. Financial and ecological management plans should be developed that weigh the loss of small, domesticated livestock through depredation up against the benefits of taking an ecosystem approach to the husbandry of small, domesticated livestock. Inclusion of such financial and ecological management plans could become the point of departure for the Western Cape Nature Conservation Board when evaluating applications for carnivore control permits. The black-backed jackal and the caracal are the products of a long period of development and co-existence with humans and are well adapted to it. They have been preying on small, domesticated livestock in Africa long before the time of the European settlers. It is impossible to control their numbers except for regional and national extermination and the fences on small, domesticated livestock production units are usually not effective as barriers to their movements. It seems to be ecologically and economically more prudent to maintain functional ecosystems on small, domesticated livestock production units and to take preventative measures to reduce the contact between such livestock and their predators. Small, domesticated livestock husbandry practices cannot be ecologically and economically sustainable if they are driven only by short-term

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financial gain. The lambs and lambing season(s) are critical components that could be protected by taking preventative measures. This sometimes requires nothing more than small adjustments in the husbandry approach being taken to handling livestock depredation. Controlling the depredation by carnivores on small, domesticated livestock cannot merely be done through attempts at controlling the population size of the carnivores that are involved. A multi-faceted approach is advised that is aimed at minimizing depredation by the black-backed jackal and caracal on small, domesticated livestock. The minimization of such depredation can be achieved by taking an ecosystem approach to the husbandry of small, domesticated livestock in combination with the use of carnivore-proof overnight and maternity pens and livestock protection or herding guard dogs. The ecological benefits of the black-backed jackal and caracal through the removal of natural herbivorous prey that reduce the grazing potential of the habitat are a bonus that should form part of the overall financial balance statement. The uncontrolled use of leg-hold (gin) traps and poisoned bait are non-specific and hazardous for humans, livestock and biodiversity conservation. The removal of some wild carnivores from an ecosystem will cause their ecological role to be taken over by other carnivores. The control of carnivores also does not reduce their numbers permanently because of natural population resilience as the mortalities that are caused by control operations simply replace those that would occur naturally. The overall conclusions are that their indiscriminate killing has not succeeded in reducing the impact of black-backed jackals and caracals on small, domesticated livestock substantially and that all such efforts have failed to date. A carefully balanced approach that includes ecosystem–based husbandry and selected preventative measures appears to be the only viable alternative. 7. Specific conclusions and recommendations 1. The black-backed jackal and the caracal have lived alongside early humans millions of years. They also were associated with and caused livestock losses to the Khoikhoi herders before the arrival of settlers from Europe in South Africa. 2. Their wide geographic distribution and rapid population recovery rate make it economically and ecologically impossible to eradicate the black-backed jackal and the caracal populations locally and regionally in South Africa.

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3. All attempts to control black-backed jackal and caracal numbers through intensive control campaigns at great cost have failed over at least the past six decades. 4. Most lethal control methods are not specific in terms of the species or individual being targeted and consequently are not options that are economically and ecologically effective, except in localized, short-term control attempts under specific conditions of ecosystem management which are aimed at reaching specific objectives. 5. Methods to prevent or limit depredation on small livestock at specific times are the only option that is viable over the long term although many of them are only partially effective. 6. The benefits and costs of an ecosystem-based husbandry approach to the production of small, domesticated livestock should be included in any husbandry programme. Such an approach will prevent overgrazing and trampling of the habitat, supply an alternative food resource to black-backed jackals and caracals in the form of healthy populations of smaller mammals, and should also contain a quantitative financial management plan as a long-term option. One objective would be to determine critical periods when short-term intensive control of carnivore numbers is likely to be ecologically and economically desirable and to measure the real financial and ecological costs and benefits of carnivore control for small, domesticated livestock production. 7. Depredation on small, domesticated livestock by habituated carnivores can be countered by increasing ecosystem functioning which would also increase livestock productivity and offset losses. Such an approach should be combined with the use of carnivore-proof overnight and maternity pens and livestock protection or herding guard dogs that will protect livestock by day and night as this has proved to be effective elsewhere. 8. As only a coyote-getter that is set by a professionally trained person has some measure of selectiveness to target a so-called “problem” carnivore when it is set around a carnivore-proof livestock enclosure at night, this device could be used to remove individual carnivores that become persistent habitual livestock killers. However, it should not remain in place for more than 10 days. 9. Methods of controlling carnivore numbers such as the use of coyote-getters can also be used to reduce the carnivore population on a livestock production unit temporarily for at most the month before the lambing season starts, provided that this is economically feasible. 10. It is recommended that short-term permits to control carnivore populations only be granted to those producers who are following an ecosystem-based approach to the husbandry of small, domesticated livestock in combination with

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effective methods to prevent depredation. Applications for carnivore control permits should also be accompanied by viable ecosystem and financial management plans. 11. It seems to be time for a unified approach in which all parties will have to yield some ground so that all may eventually benefit from developing a new and holistic strategy. In doing so, limited agendas have to be traded for broader visions in a spirit of true cooperation. 12. The Western Cape Nature Conservation Board cannot adopt an approach that will lead to the degradation of biodiversity and is only aimed at the economic benefits of livestock production because that is not their mandate.

13. It is suggested that the Western Cape Nature Conservation Board facilitate the following research projects: - The ecological and economic advantages of taking an ecosystem approach to small, domesticated livestock husbandry. - The role of selected conservation areas in the Western Cape province as “breeding grounds” for black-backed jackals and caracals on adjacent small, domesticated livestock production units. - The effectiveness of using livestock protection or herding guard dogs to prevent depredation on small, domesticated livestock by the black-backed jackal and caracal. - The effectiveness of using carnivore-proof overnight and maternity pens in reducing or preventing depredation by the black-backed jackal and caracal on small, domesticated livestock. - The effectiveness of livestock fences to curtail the movements of the black-backed jackal and caracal. - The distances that young black-backed jackals and caracals disperse from their natal areas. - Population recovery rates of black-backed jackals and caracals following the control of their numbers on small. Domesticated livestock production units. Complete bibliography Andelt, W F 1992. Effectiveness of livestock guarding dogs for reducing predation on domestic sheep. Wildlife Society Bulletin 20(1): 55 – 62. Anon 1986. Proceedings of a colloquium on the use/abuse of strychnine. Onderstepoort: The Wildlife Group of the South African Veterinary Association, 15 February. Anon 1992. Oranjejag carnivore control statistics. Orange Free State: Oranjejag. Anon 2008. Researchers say more gray wolves mean more pronghorn antelope in the Yellowstone area. Associated Press, 3 March.

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Anon 2010. Anatolian shepherd livestock guarding dog project. Progress Report: Cheetah Outreach, December 2010. Anon 2011. The black-backed jackal, http://en.wikipedia.org/wiki?Black-backedJackal Avenant, N L 1993. The caracal, Felis caracal Schreber, 1776, as predator in the West Coast National Park. MSc dissertation. Stellenbosch: University of Stellenbosch. Avenant, N L and J A J Nel 1997. Prey use by four syntopic carnivores in a Strandveld ecosystem. South African Journal of Wildlife Research 27(3): 86 – 93. Avenant, N L and J A J Nel 1998. Home-range use, activity, and density of caracal in relation to prey density. African Journal of Ecology 36: 347 – 359. Avenant, N L and J A J Nel 2002. Among habitat variation in prey availability and use by Felis caracal. Mammalian Biology 67: 18 – 33. Berger, J 2007. Fear, human shields and the redistribution of prey and predators in protected areas. Biology Letters 3: 620 – 623. Berger, J, P B Stacey, L Bellis and M P Johnson 2008. A mammalian predator-prey imbalance: grizzly bear and wolf extinction affect avian Neotropical migrants. Ecological Applications 11(4): 947 – 960. Berger, K M 2006. Carnivore-livestock conflicts: effects of subsidized predator control and economic correlates on the sheep industry. Conservation Biology 20(3): 751 – 761. Bernard, R T F and C T Stuart 1987. Reproduction of the caracal Felis caracal from the Cape Province of South Africa. South African Journal of Zoology 22: 177 – 182. Bester, J L 1982. Die gedragsekologie en bestuur van die silwervos Vulpes chama (A Smith) met spesiale verwysing na die Oranje Vrystaat. MSc dissertation. Bloemfontein: University of the Orange Free State. Bingham, J and G K Purchase 2002. Reproduction in the jackals Canis adustus Sundevall, 1846 and Canis mesomelas Schreber, 1778 (Carnivora: Canidae), in Zimbabwe. African Zoology 37(1): 21 – 26. Bingham, J and G K Purchase 2002. Age determination in jackals Canis adustus Sundevall, 1846 and Canis mesomelas Schreber, 1778 (Carnivora: Canidae)

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with reference to the age structure and breeding patterns of jackal populations in Zimbabwe. African Zoology 38(1): 153 – 160. Blejwas, K M, B J Sacks, M M Jaeger and R McCullough 2002. The effectiveness of selective removal of breeding coyotes in reducing sheep predation. Journal of Wildlife Management 66(2): 451 – 462. Brand, D J 1989. The control of caracal (Felis caracal) and baboons (Papio ursinus) in the Cape Province with the help of mechanical means. MSc dissertation. Stellenbosch: University of Stellenbosch. Bothma, J du P 1965. Random observations on the food habits of certain Carnivora in South Africa. Fauna and Flora 16: 16 – 22. Bothma, J du P 1971. Control and ecology of the black-backed jackal Canis mesomelas in the Transvaal. Zoologica Africana 6(2): 187 – 189. Bothma, J du P 1971. Food of Canis mesomelas in South Africa. Zoologica Africana 6(2): 195 – 203. Bothma, J du P 1971. Notes on movement by the black-backed jackal and the aardwolf in the western Transvaal. Zoologica Africana 6(2): 205 – 207. Bothma, J du P 1993. Probleemdiere slagoffers van naamsverskille. Landbouweekblad, 24 December: 18 – 21. Bothma, J du P 1998. Carnivore ecology in arid lands. Berlin: Springer. Bothma, J du P 2010. Jackals. Game and Hunt 16(2): 6 - 9. Bothma, J du P. 2010. Predators: general principles. In J du P Bothma and J G du Toit (Eds), Game ranch management, fifth edition. Pretoria: Van Schaik, pp 289 – 295. Bothma, J du P 2010. Aerial counts of wild animals. In J du P Bothma and J G du Toit (Eds), Game ranch management, fifth edition. Pretoria: Van Schaik, pp 467 – 476. Bothma, J du P and E A N le Riche 1984. Aspects of the ecology and the behaviour of the leopard Panthera pardus in the Kalahari Desert. Supplement to Koedoe 1984: 259 - 279. Bothma, J du P and E A N le Riche 1986. Prey preference and hunting efficiency of the Kalahari Desert leopard. In S D Miller and D D Everett (Eds), Cats of the world: biology, conservation and management. Washington, DC: National Wildlife Federation, pp 389 – 414.

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Bothma, J du P and E A N le Riche 1994. Scat analysis and aspects of defecation in northern Cape leopards. South African Journal of Wildlife Research 24(1&2): 21 – 25. Bothma, J du P and E A N le Riche 1994. Range use by an adult male caracal in the southern Kalahari. Koedoe 37: 105 – 108. Bothma, J du P, J A J Nel and A Macdonald 1984. Food niche separation between four sympatric Namib Desert carnivores. Journal of Zoology (London) 202: 327 – 340. Bothma, J du P and C Walker 1999. Larger carnivores of the African savannas. Pretoria: Van Schaik, pp 116 – 130. Brain, C K 1993. Swartkrans: a cave’s chronicle of early man. Transvaal Museum Monograph 8: 1 – 270. Bromley, C and E M Gese 2001. Surgical sterilization as a method of reducing coyote predation on domestic sheep. Journal of Wildlife Management 65(3): 510 – 519. Burns, R J and G E Connolly 1995. Assessment of potential toxicity of Compound 1080 from livestock protection collars to canines and scavenging birds. International Biodeterioration and Biodegradation 1995: 161 – 167. Connolly, G E 1978. Predator control and coyote populations: a review of simulation models. Coyotes. New York: Academic Press. Carbone, C, A Teacher and J M Rowcliffe 2007. The costs of carnivory. PLoS Biology 5(2): 363 - 368. Carlson, D A and E M Gese 2009. Integrity of mating behaviours and seasonal reproduction in coyotes (Canis latrans) following treatment with estradiol benzoate. Animal Reproduction Science 117: 322 – 330. Connor, M M, M R Ebinger and F F Knowlton 2008. Evaluating coyote management strategies using a spatially explicit, individual-based, socially structured population model. Ecological Modelling 219: 234 – 247. Conradie, B, J Piesse and C Thirtle 2009. What is the appropriate level of aggregation for productivity indices? Comparing district, regional and national measures. Agrekon 48(1): 9 – 20.

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De Vos, V 1970. Pseudo pregnancy in the black-backed jackal (Canis mesomelas Schreber). Journal of the South African Veterinary and Medical Association 40(4): 381 – 383. Dolbeer, R A, N R Holler and D W Hawthorne 1996. Identification and control of wildlife damage. In T A Bookhout (Ed), Research and management techniques for wildlife and habitats, fifth revised edition. Bethesda: The Wildlife Society, pp 474 – 506. Dragesco-Joffé, A 1993. La vie sauvage au Sahara. Paris: Delachaux et Niestlé. Du Plessis, S S 1972. Ecology of the blesbok with special reference to productivity. Wildlife Monograph 30: 1 – 70. Dyer, J 1989.Bekendstelling van kleinvee-beskermingshalsband (livestock protection collar). Worcester: The Livestock Protection Co. Eccard, J A, R B Walther and S J Milton 2000. How livestock grazing affects vegetation structures and small mammal distribution in the semi-arid Karoo. Journal of Arid Environments 46: 103 – 106. Eloff, F C 1973. Lion predation in the Kalahari Gemsbok National Park. Journal of the southern African Wildlife Management Association 3(2): 59 – 63. Fairall, N 1968. The reproductive seasons of some mammals in the Kruger National Park. Zoologica Africana 3: 189 – 210. Ferguson, J W H 1978. Social interactions of black-backed jackals Canis mesomelas in the Kalahari Gemsbok National Park. Koedoe 21: 151 – 162. Ferguson, J W H, J A J Nel and M J de Wet 1983. Social organization and movement patterns of black-backed jackals Canis mesomelas in South Africa. Journal of Zoology (London) 199: 487 – 502. Ferguson, J W H, J S Galpin and M J de Wet 1988. Factors affecting the activity patterns of black-backed jackals Canis mesomelas. Journal of Zoology (London) 214: 55 – 59. Gehring, T M, K C verCauteren and J Landry 2010. Livestock protection dogs in the 21st Century: is an ancient tool relevant to modern conservation challenges? BioScience 60(4): 299 – 308. Goldenberg, M, F Goldenberg, S M Funk, J Henschel and E Millesi 2010. Diet composition of black-backed jackals, Canis mesomelas in the Namib Desert. Folia Zoologica 59(2): 93 – 101.

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Grafton, R N 1965. Food of the black-backed jackal: a preliminary report. Zoologica Africana 1(1): 41 – 53. Green, J S, R A Woodruff and R Horman 1984. Livestock guarding dogs and predator control. Rangelands 6(2): 73 – 76. Grobler, J H 1981. Feeding behaviour of the caracal Felis caracal in the Mountain Zebra National Park. South African Journal of Zoology 16: 259 – 262. Grobler, J H 1982. Growth of a male caracal kitten Felis caracal in the Mountain Zebra National Park. Koedoe 25: 117 – 119. Grobler, J H, A Hall-Martin and C Walker 1984. Predators of southern Africa. Johannesburg: Macmillan. Gusset, M, M J Swarner, L Mponwane, K Ketetile and J W Mc Nutt 2009. Human-wildlife conflict in northern Botswana: livestock predation by endangered African wild dog Lycaon pictus and other carnivores. Oryx 43(1): 67 – 72. Harrington, J T and M R Conover 2007. Does removing coyotes for livestock protection benefit free-ranging ungulates? Journal of Wildlife Management 71(5): 1555 – 1560. Hansen, I, T Staaland and A Ringsø 2002. Patrolling with livestock guard dogs: a potential method to reduce predation on sheep. Acta Agriculturae Scandanavica, Section A, Animal Science 52: 43 – 48. Hawley, J E, T M Gehring, R N Schulz, S T Rossler and A P Wydeven 2009. Assessment of shock collars as nonlethal management for wolves in Wisconsin. Journal of Wildlife Management 73(4): 518 – 525. Hayward, M W and G J Hayward 2010. Potential amplification of territorial advertisement markings by black-backed jackals (Canis mesomelas). Behaviour 147: 979 – 992. Herfindal, I, J D C Linnell, P F Moa, J Odden, L B Austmo and R Andersen 2005. Does recreational hunting reduce losses on domestic sheep? Journal of Wildlife Management 69(3): 1034 – 1042. Hiscocks, K and M R Perrin 1988. Home range movements of black-backed jackals at Cape Cross Seal Reserve, Namibia. South African Journal of Wildlife Research 18(3): 97 – 100. Kamler, J F, N F Jacobsen and D W Macdonald 2008. Efficiency and safety of Soft Catch traps for capturing black0backed jackals and excluding non-target species. South Africa Journal of Wildlife Research 38(2): 113 – 116.

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