A dissertation submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg in fulfilment of the requirements for the degree of Master of Science, 2017 / The African wild dog (Lycaon pictus) is one of South Africa’s most endangered carnivores. The species has suffered massive range shrinkages in the past few decades and population numbers have dropped significantly. Along with other factors responsible for its decline, one of the most notable threats to the African wild dog is conflict with humans. These carnivores are often persecuted by farmers for their alleged depredation of livestock and captive-bred game species, although doubt exists as to whether wild dogs are the avid depredators as suggested. My research therefore aimed to investigate the conflict between people and African wild dog, focussing on livestock depredation. Due to heterogeneous farming landscapes, the history and location of protected areas and the endangered status of the African wild dog, South Africa provides many opportunities to study this particular type of human-carnivore conflict. Firstly, I conducted a meta-analysis of human-carnivore conflict using published literature about African wild dog depredation of livestock and game and compared these to other African carnivores as well as non-African carnivores. Results indicated that African wild dog were less avid depredators than other African species such as lion (Panthera leo) and spotted hyena (Crocuta crocuta). Also evident was that high carnivore and livestock densities, coupled with poor communities with poor livestock husbandry practices, make people and carnivores in developing regions more vulnerable to human-carnivore conflicts. Secondly, I assessed actual African wild dog occurrence in relation to the location of farms, livestock density and several other anthropogenic and natural landscape features. This was achieved using GPS data from four collared African wild dog individuals from packs residing in the northeastern part of South Africa and resource selection functions. Results from these analyses suggested that, whilst African wild dog may occur in close proximity to farms, they established home ranges in areas of low livestock density and few farms, indicating predictive avoidance of areas where mortality may occur. Major roads were highlighted as a vulnerability for the African wild dog, whilst nature reserves and vegetation were also important predictors of wild dog occurrence. Other anthropogenic and natural landscape features varied in importance in determining wild dog occurrence. Knowledge about how the African wild dog selects its resources will enable us to identify vulnerabilities for these carnivores as well as areas where they are likely to occur, aiding in conservation planning. Though African wild dog have historically been reported to kill livestock such goats and cattle, my study seems to indicate that these carnivores are not avid stock-killers. Given the precarious survival status of the African wild dog and the food security needs of people in a developing region strongly suggests the need for cooperation of farmers and the education of communities to aid the recovery of this uniquely African carnivore. / XL2018

The African wild dog, second most endangered carnivore in Africa, has a well-developed, highly cooperative pack system. The usual structure of a pack consists of a dominant breeding pair, the alpha male and female, several subordinates, non-breeding adults and dependent offspring. Domestic dog microsatellites were used to study the parentage in three packs and confirm that more than one dog, including the subordinate males, can sire pups within a litter as previously suggested. The study was performed on two isolated populations of wild dogs in the North West Province of South Africa. In Madikwe Game Reserve, skin samples from 47 dogs were obtained by means of biopsy darts (adults) and skin slivers taken from the ear (subadults) and stored in absolute ethanol. In Pilanesberg National Park, blood samples from 18 captured dogs were collected in EDTA blood tubes. The wild dogs were photographed and individually identified according to coat patterns. Behavioural data to determine ranking were collected from all three packs. DNA was extracted from collected samples using proteinase-K digestion followed by isolation of DNA with phenol/chloroform/isoamyl alcohol. A total of 16 microsatellite loci that consistently amplified and appeared to be polymorphic in wild dogs, were used. Polymerase Chain Reaction (PCR) was performed using two panels of microsatellite loci in multiplex reactions. An amount of 1 µl of PCR product was loaded on to the 3130 XL Genetic Analyser with Genescan 500 LIZ (Applied Biosystems) size standard and analysed using STRand (Board of Regents, University of California) software program. CERVUS 2.0 software was used to calculate allele frequencies, expected and observed heterozygosity, frequency of null alleles, polymorphic information content and exclusion probabilities for parentage assignment. Parentage verification was also performed manually. The parentage analysis revealed that at least one pup was not sired by the alpha male in each of the five litters studied. Although previous studies suggested that the alpha male sires the majority of offspring in the pack, our results confirm that subordinate males commonly sire pups with the alpha female if and when the opportunity arises. This is possibly a mechanism to decrease the effects of inbreeding. / Dissertation (MSc (Veterinary Sciences))--University of Pretoria, 2006. / Production Animal Studies / unrestricted

Doctor of Philosophy / The African wild dog (Lycaon pictus) is one of Africa’s most endangered carnivores. Previous research into this species has focussed on the largest extant populations in Africa. However, there are a large number of relatively small populations (20 to 50 dogs) distributed across Africa, which represent an important component of the diversity of the species and its remaining habitat. This study investigated the status of a small population of wild dogs in the Lower Zambezi area in Zambia. Objectives focussed on assessing population dynamics and identifying causes of decline. Research was carried out over a broad range of topics in an effort to provide comprehensive information for conservation management of the population. The scope of the project was divided into five sections: 1) Demography and pack dynamics were assessed to identify the structure and status of the population, and the main causes of mortality. 2) An assessment of habitat types and related ecological factors was carried out to determine wild dog habitat utilisation in relation to vegetation type, prey densities and hunting success in each area. 3) The effects of interpredator competition on wild dog population dynamics was investigated, specifically, the effects of lions (Panthera leo) and spotted hyaenas (Crocuta crocuta). 4) Genetic analyses were carried out to assess the historic and contemporary genetic variability of the population, and to define patterns of geographic structuring and population differentiation. 5) Results were combined to assess the viability of the population and recommend conservation management strategies. Snaring was identified as the most important cause of adult mortality, and a threat to wild dog population persistence. Inbreeding avoidance led to the emigration of adult males and females from the area and appeared to be a substantial contributor to population decline. Limited mate selection corresponded with neither sex displaying philopatry and large dispersal distances effectively removed adults from the population. This result has important implications for the management of small populations, whereby lack of mate choice may increase dispersal distances and thereby increase edge effects on populations, regardless of home range sizes. Home range sizes were related to den locations in remote areas of the Zambian Escarpment, which was used as a breeding refuge area. The Zambezi River and Zambian Escarpment appeared to be effective barriers to wild dog home range movements. The study area contained a diversity of habitats on the alluvial terraces of the river valley floor. There was a high density of impala (Aeypceros melampus), which formed the main prey base for the wild dog population. Studies of other populations have found that wild dogs often avoided areas with high competing predator densities, which corresponded with high prey density areas. In contrast to those findings, the Lower Zambezi wild dog population showed a strong preference for high prey density areas. This population also showed only temporal avoidance of high lion density areas. Low lion density areas were preferred during breeding periods, while moderate to high lion density areas were preferred during non-breeding periods. Direct predation of adult wild dogs by lion and spotted hyaenas was rare. Kleptoparasitism of wild dog kills by either competing predator species was also rare. Predator competition was not considered to be an important determinant of population decline. The Lower Zambezi population suffered from a loss of heterozygosity, low allelic richness, and there was significant evidence of a recent population bottleneck. The population did not contain any new mtDNA haplotypes, nor any unique alleles on the commonly used microsatellite loci, but was differentiated from African wild dog populations in other regions. There was evidence of historical and recent gene flow between the Lower Zambezi and the neighbouring southern African populations of Hwange and Okavango. This was the first study to show a loss of genetic variability in a free-ranging African wild dog population. Although more immediate anthropogenic and demographic factors were the critical determinants of population decline, the loss of genetic variability has important implications for the conservation of the remaining small and fragmented wild dog populations in Africa. Results showed that due to its small size the population is likely to have suffered from inverse density dependence and Allee effects on dispersal and reproductive success. Management recommendations focussed on mitigating anthropogenic causes of mortality, and improving connectivity with a larger, potential source population to increase the probability of successful dispersal and to restore genetic diversity. The high density prey base, small home range sizes and low levels of interpredator competition detected in this study suggest that the area has the capacity to support a large and potentially viable population of wild dogs if appropriate management strategies are implemented.

The obligate cooperative nature of African wild dogs (Lycaon pictus), unique among social canids, is thought to be driven by high levels of interspecific competition and intra-guild predation with other large socially-organised predators. Research exploring how wild dogs maintain social bonds through social communication, while avoiding detection from potential competitors, is therefore vital for understanding this species. While olfactory and vocal communications are well represented in the literature concerning L. pictus, these channels of communication pose significant risks to wild dog survival, as they are inherently susceptible to eavesdropping by unintended receivers. In comparison, display communication, which requires visual contact between the signaller and receiver, poses comparatively less risk of attracting the attention of eavesdroppers. In spite of this, few studies have explored the use of display communication in wild dogs, leaving its potential significance in maintaining social bonds within packs unexplored. Using video analysis, I investigate how display communication and subsequent social interaction are affected by several pressures of the captive environment. Captivity did not appear to affect the presence of many social display types also used by free-ranging wild dogs. However, sexual behaviour was absent from the study groups, likely due to the use of contraception and pack sex composition. The effects of pack sex composition (single-sex versus mixed-sex) revealed that while the frequencies and durations of many social behaviours were not affected by sex composition, the distribution of social interactions did differ depending on group structure. Here, a highly related, single-sex group was more stable than a highly related, mixed-sex group of the same size. Differences in the captive management strategies, specifically enclosure size and feeding regime, of the two packs, may explain the behavioural differences observed. The death of a pack member permitted investigation of the effects of death on social interactions. While many of the social interactions were unaffected by the death, behaviours relating to the formation and maintenance of social dominance and social hierarchy increased after the dog’s death. The distribution of social interactions was non-random, suggesting that individuals were reorganising the social structure of the pack during this period. Finally, a small study into the effects of simultaneously added enrichments in the pack (post death) revealed that enrichment reduced the frequencies of dominance behaviours and allowed for a more even distribution of social interaction within the pack. This demonstrates how enrichment may potentially be used to reduce aggression within captive animals. Overall, this research reveals that display-based communication is important for the maintenance of sociality in captive African wild dogs. To better understand this endangered species, future studies in free-ranging populations should include this channel of communication.

Doctor of Philosophy / The African wild dog (Lycaon pictus) is one of Africa’s most endangered carnivores. Previous research into this species has focussed on the largest extant populations in Africa. However, there are a large number of relatively small populations (20 to 50 dogs) distributed across Africa, which represent an important component of the diversity of the species and its remaining habitat. This study investigated the status of a small population of wild dogs in the Lower Zambezi area in Zambia. Objectives focussed on assessing population dynamics and identifying causes of decline. Research was carried out over a broad range of topics in an effort to provide comprehensive information for conservation management of the population. The scope of the project was divided into five sections: 1) Demography and pack dynamics were assessed to identify the structure and status of the population, and the main causes of mortality. 2) An assessment of habitat types and related ecological factors was carried out to determine wild dog habitat utilisation in relation to vegetation type, prey densities and hunting success in each area. 3) The effects of interpredator competition on wild dog population dynamics was investigated, specifically, the effects of lions (Panthera leo) and spotted hyaenas (Crocuta crocuta). 4) Genetic analyses were carried out to assess the historic and contemporary genetic variability of the population, and to define patterns of geographic structuring and population differentiation. 5) Results were combined to assess the viability of the population and recommend conservation management strategies. Snaring was identified as the most important cause of adult mortality, and a threat to wild dog population persistence. Inbreeding avoidance led to the emigration of adult males and females from the area and appeared to be a substantial contributor to population decline. Limited mate selection corresponded with neither sex displaying philopatry and large dispersal distances effectively removed adults from the population. This result has important implications for the management of small populations, whereby lack of mate choice may increase dispersal distances and thereby increase edge effects on populations, regardless of home range sizes. Home range sizes were related to den locations in remote areas of the Zambian Escarpment, which was used as a breeding refuge area. The Zambezi River and Zambian Escarpment appeared to be effective barriers to wild dog home range movements. The study area contained a diversity of habitats on the alluvial terraces of the river valley floor. There was a high density of impala (Aeypceros melampus), which formed the main prey base for the wild dog population. Studies of other populations have found that wild dogs often avoided areas with high competing predator densities, which corresponded with high prey density areas. In contrast to those findings, the Lower Zambezi wild dog population showed a strong preference for high prey density areas. This population also showed only temporal avoidance of high lion density areas. Low lion density areas were preferred during breeding periods, while moderate to high lion density areas were preferred during non-breeding periods. Direct predation of adult wild dogs by lion and spotted hyaenas was rare. Kleptoparasitism of wild dog kills by either competing predator species was also rare. Predator competition was not considered to be an important determinant of population decline. The Lower Zambezi population suffered from a loss of heterozygosity, low allelic richness, and there was significant evidence of a recent population bottleneck. The population did not contain any new mtDNA haplotypes, nor any unique alleles on the commonly used microsatellite loci, but was differentiated from African wild dog populations in other regions. There was evidence of historical and recent gene flow between the Lower Zambezi and the neighbouring southern African populations of Hwange and Okavango. This was the first study to show a loss of genetic variability in a free-ranging African wild dog population. Although more immediate anthropogenic and demographic factors were the critical determinants of population decline, the loss of genetic variability has important implications for the conservation of the remaining small and fragmented wild dog populations in Africa. Results showed that due to its small size the population is likely to have suffered from inverse density dependence and Allee effects on dispersal and reproductive success. Management recommendations focussed on mitigating anthropogenic causes of mortality, and improving connectivity with a larger, potential source population to increase the probability of successful dispersal and to restore genetic diversity. The high density prey base, small home range sizes and low levels of interpredator competition detected in this study suggest that the area has the capacity to support a large and potentially viable population of wild dogs if appropriate management strategies are implemented.

Population estimation is an important task in all wildlife conservation. Such estimations are often difficult in low-density species such as big carnivores. The painted dog (Lycaon pictus) is an endangered species and the first aim of IUCNs action plan is to assess the size and the distribution of the remaining population. This study is the first step towards a quantitative method to distinguish painted dog footprints from footprints of feral domestic dogs and hyaenas. Footprint photographs were collected and digitally processed and total pad area and angles between the digits and backpad of the paw were measured. Both the pad area and the angles show a statistically significant difference between the species. However, further analysis shows that there is no significant difference in pad area between painted dog females and domestic dog males. Size of the pads alone is therefore not suitable as a measure to determine the species from an unknown footprint. The angles between backpad and digits seem to be more suitable to distinguish between species. Therefore, a combination of pad size and the angle between backpad and digits might be useful to estimate the species from an unknown footprint in the field.

The African wild dog (Lycaon pictus) was reintroduced into Hluhluwe Umfolozi Park (HUP) in 1980/81 after an absence of over 50 years. A 'hands off' management approach was applied. Although the reintroduction is regarded as successful, the population has not increased significantly and still only consists of a single pack. Various aspects which may affect this population were investigated to compile an active management strategy. Home range analyses identified extensive use of a home range which covered only 22.7 % of HUP and was situated entirely in Hluhluwe Game Reserve. The extensive movements of the dogs within their home range were primarily dictated by the movements and distribution of their prey species, especially nyala (Tragelaphus angasu) and impala (Aepyceros melampus). Forest habitats were preferred, but all available habitat types were utilised extensively by the wild dogs. Space and habitat were both considered to have no limiting effect on this population and HUP has sufficient space and suitable habitat to support a larger population of L. pictus. A number of potentially pathogenic antigens were identified in the population, indicating that it had been exposed to these at some time. Domestic dogs in the areas surrounding HUP were identified as the most likely source of these diseases which pose a threat to the wild dogs. Some genetic considerations are discussed in terms of their effect on the population status and management of the wild dogs. A number of management proposals based on the results of the project were formulated. It is suggested that an active management approach be adopted for the population. This includes managing it as part of a metapopulation which primarily involves the exchange of genetic material among similar wild dog populations on a regular basis. The immediate supplementation of the population with new genetic material is proposed. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 1996.

The small population size of wild dog Lycaon pictus (10) in Hluhluwe-Umfolozi Park (HUP) and the decline in their numbers since 1992, has caused concern for their survival and consideration of further introductions. In the light of many failed wild dog relocation and reintroduction programmes, this study contributes towards an understanding of the ecology of the HUP wild dog pack. Wild dog prey preference was determined from scat analysis and personal observations, and their potential impact on the primary prey species was modelled. The choice of physical habitat features by wild dog and their ranging behaviour within the Park were correlated with the distribution of their primary prey and other predators. To determine the susceptibility of prey to predation in three reserves with different predator diversities and densities, prey vigilance and prey response to playback recordings of predator calls were recorded. The results showed that wild dog preference for females, adult nyala Tragelaphus angasi and juvenile impala Aepyceros melampus, was a function of prey abundance, profitability calculated using a diet choice model, and ease of capture. Based on the overall lack of association of wild dog and their primary prey species and predators, and the overall lack of similarity of wild dog and prey choice of physical habitat features, predator presence was the most important determinant of wild dog ranging behaviour. Prey vigilance differed significantly between reserves and was inversely correlated with predator density. Prey response to predator calls did not differ significantly between reserves but prey did, however, react sooner to those calls unfamiliar to them. Nyala were more vigilant and responded sooner to playbacks than impala suggesting that nyala may experience greater levels of predation pressure. There was no evidence to suggest that the prey preference, habitat preference and ranging behaviour of the wild dog were influenced by the susceptibility of prey to predation. Models of prey population dynamics determined that although the introduction of an additional wild dog pack would result in a reduction of current prey population growth rates and an increase in prey population extinction probabilities, their predicted impact would be slight. Since emigration and population viability were identified as the primary causes of the HUP wild dog population decline, the introduction of two groups of wild dog individuals into Hluhluwe was suggested to boost population numbers and stimulate breeding and dispersal within the Park. The importance of future monitoring and proactive management was stressed to ensure the survival of this valuable species in the Park. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 1996.

The African wild dog Lycaon pictus is Africa’s second most endangered carnivore. Only 14 out of 39 countries in Africa still have wild dogs present. This makes the populations of wild dogs in South Africa very valuable with respect to the entire species. Kruger National Park (Kruger) has the only self-sustaining and viable population of wild dogs in South Africa, making Kruger the core area of conservation for South African wild dogs. It is of vital importance to know the numbers of wild dogs present in Kruger. In chapter 2 of this dissertation I monitored and gathered demographic information from as many southern Kruger wild dog packs and individuals as possible over a three month period. I used real time text messaging to collect the information. A wild dog hotline number was used for tourists to contact immediately after they sighted a pack, noting location, time and number of wild dogs sighted. This new technique resulted in more than 300 reported wild dog sightings in three months enabling a count of individuals and packs. This also created an opportunity to take identification photographs and to collect DNA samples. In 1997 it was decided to establish and manage several small wild dog populations in various geographically isolated reserves in South Africa as one large managed metapopulation. In order to simulate the natural dispersal patterns of wild dogs, individuals are translocated between the managed metapopulation reserves, imitating natural gene flow and hopefully preventing inbreeding. To date, all decisions have been made using demographic data only. This in time is likely to result in a loss of genetic diversity and subsequent inbreeding. The aim of chapter 3 was to obtain genetic information from wild dogs in the managed metapopulation and Kruger (chapter 2) to provide a basis for sound population management including monitoring of inbreeding and maintaining levels of genetic diversity similar to those found in large self-sustaining populations (such as Kruger). This study included both mitochondrial DNA (mtDNA) and nuclear microsatellite loci to determine the genetic structure of South Africa’s wild dogs specifically with regards to genetic diversity, population structure and relatedness. The results showed a difference in historical and recent diversity between the managed metapopulation and Kruger. Two genetic clusters were evident in South Africa, however one was due to wild dogs from Botswana being translocated into the managed metapopulation. After the Botswana influence was removed from the analysis, three genetic clusters were observed in the South African wild dogs. These three genetic clusters comprise too few wild dogs to manage them as separate units. Relatedness between and within populations, reserves and packs were estimated and can in future be used to guide translocations of wild dogs to maximise their genetic variability. It is suggested that due to the low numbers, and historical and recent trends in genetic structure of South Africa’s wild dogs, they should be managed as one unit, allowing movements to and from neighbouring countries. All translocations should follow an isolation-by-distance pattern. / Dissertation (MSc)--University of Pretoria, 2010. / Animal and Wildlife Sciences / unrestricted

The African wild dog (Lycaon pictus) is the second most endangered carnivore in Africa. Although several high-profile endangered species are imperiled due to poor fertility, inadequate genetic diversity, and a reliance on specific niches, the wild dog is threatened by decreasing land space and human hunting practices. Wild dogs are highly fertile with an average litter size of nine at De Wildt Cheetah and Wildlife Centre. Wild dog males have 3 million to 900 million sperm per ejaculate and 45-75% progressive motility during the breeding season. Wild dogs released into small nature reserves in South Africa experience increased survival rates due to sizeable litters, abundant prey, and increased hunting success along fence lines. Furthermore, the current demand for captive-bred wild dogs is low due to lack of demand by overseas zoos and the inability of nature reserves to accommodate more dogs. Long-acting GnRH analogues have been used for fertility control in many wildlife species. However, dosing and efficacy differ among species and individual animals. This study assessed the efficacy of the GnRH analogue, deslorelin, on reproductive parameters of male African wild dogs. Seasonal effects on reproduction were also evaluated. Captive male African wild dogs housed at the De Wildt Cheetah and Wildlife Centre were administered either a 4.7 mg deslorelin implant (Suprelorin,® Peptech Animal Health (Pty) Ltd, Sydney, Australia; n = 10), an experimental 9.4 mg deslorelin injection (n = 11) or a placebo injection (n = 6). Treatment was administered during the non-breeding season (Month 0), and dogs were assessed at Months 3, 5, 6, and 7. Reproductive parameters evaluated before and after treatment included: serum testosterone, testicular and prostatic volume, and semen quality. Serum testosterone was assessed with a previously validated double antibody DSL testosterone radioimmunoassay kit (Diagnostic Systems Laboratories, Inc, Webster, TX). Testicular volume was calculated from testicular dimensions measured with a calliper and prostatic volume from dimensions obtained by trans-cutaneous ultrasound. Data were analyzed with ANOVA. Although the 4.7 mg deslorelin implant was safe for use in male wild dogs, there was wide variation in efficacy among dogs. The serum testosterone of implant dogs did not decrease to baseline after treatment and only half the dogs administered an implant became azoospermic post-treatment. The experimental long-acting deslorelin injection was ineffective for contraception of male African wild dogs. All three groups of dogs experienced an improvement in reproductive parameters during the months of February through May, the rainy season in northern South Africa and the period during which female African wild dogs enter oestrus, suggesting that a breeding season not only exists in the female African dog but also in the male. Testis and prostatic volume increased, serum testosterone concentrations and semen quality improved during that time of year. Further studies of deslorelin in male wild dogs are warranted to determine the appropriate dose, pay-out pattern, delivery method, and season of delivery necessary for adequate contraception in this species. / Dissertation (MSc)--University of Pretoria, 2008. / Production Animal Studies / unrestricted