Conservation implications of prey responses to wild dogs Lycaon pictus during the denning season on wildlife ranches

The spread of game ranching in southern Africa provides opportunities for the reestablishment of populations of endangered wild dogs extirpated by livestock ranchers. However, this potential has not been realized, partly because of negative rancher perceptions. Some ranchers believe that wild dogs impart costs by killing wildlife that could be utilized consumptively. Others complain that wild dogs make ungulates 'skittish' and cause local reductions in prey densities while denning. We compared the skittishness and density of prey species inside and outside the denning home ranges of nine wild dog packs in Zimbabwe. Wild dogs had no impact on prey skittishness, but prey species did occur at lower densities inside denning home ranges. In some scenarios, and particularly on fenced game ranches, wild dogs could cause prey population declines during denning. On small game ranches, the use of fences as a tool by wild dogs during hunting can increase the proportion of large prey species in their diet by up to 11 times, and thus increase the minimum area required to support that diet. In addition, game fencing is typically permeable to wild dogs but not their prey, preventing the recovery of prey populations through the natural influx of prey animals into the denning area following departure of the dogs. Wild dogs could thus impose significant financial costs to game ranchers hosting denning packs. Our findings emphasize the importance of promoting the formation of conservancies, where neighbouring landowners remove boundary fences to create larger contiguous wildlife areas.     

The Effect of Simulated African Wild Dog Presence on Anti‐predator Behaviour of Kudu and Impala

In this study, we examined the behavioural, temporal and spatial effects of simulated African wild dog (Lycaon pictus) presence on its two main prey species: kudu (Tragelaphus strepsiceros) and impala (Aepyceros melampus). We spread African wild dog faeces around waterholes and played African wild dog sounds at different intervals to mimic immediate and non‐immediate predation pressure. We looked at anti‐predator behaviour at both a herd and individual level and distinguished between high‐quality (detracts from all other activities), high‐cost vigilance and low‐quality (used to monitor the surrounding in spare time), low‐cost vigilance to determine costs involved. We found that simulated African wild dog presence had little effect on anti‐predator behaviour of their free‐ranging prey. Only when immediate predation risk was mimicked did kudu invest in (additional) high‐quality vigilance, whereas impala showed no response. Regardless of direct cues of African wild dog presence, behavioural adjustments to reduce predation risk were primarily based on environmental factors such as time of the day and broad‐scale habitat structure. Predators have been shown to utilize waterholes to hunt, and prey species are therefore likely to maximize anti‐predator behaviour in this high‐risk environment based on environmental variables affecting predation risk, the main predator within the system, and water requirements, leaving little flexibility to respond to (simulated) African wild dog presence.     

Demographic and functional responses of wild dogs to poison baiting

Lethal control of wild dogs – that is Dingo (Canis lupus dingo) and Dingo/Dog (Canis lupus familiaris) hybrids – to reduce livestock predation in Australian rangelands is claimed to cause continental‐scale impacts on biodiversity. Although top predator populations may recover numerically after baiting, they are predicted to be functionally different and incapable of fulfilling critical ecological roles. This study reports the impact of baiting programmes on wild dog abundance, age structures and the prey of wild dogs during large‐scale manipulative experiments. Wild dog relative abundance almost always decreased after baiting, but reductions were variable and short‐lived unless the prior baiting programme was particularly effective or there were follow‐up baiting programmes within a few months. However, age structures of wild dogs in baited and nil‐treatment areas were demonstrably different, and prey populations did diverge relative to nil‐treatment areas. Re‐analysed observations of wild dogs preying on kangaroos from a separate study show that successful chases that result in attacks of kangaroos by wild dogs occurred when mean wild dog ages were higher and mean group size was larger. It is likely that the impact of lethal control on wild dog numbers, group sizes and age structures compromise their ability to handle large difficult‐to‐catch prey. Under certain circumstances, these changes sometimes lead to increased calf loss (Bos indicus/B. taurus genotypes) and kangaroo numbers. Rangeland beef producers could consider controlling wild dogs in high‐risk periods when predation is more likely and avoid baiting at other times.     

Characteristics of African wild dog natal dens selected under different interspecific predation pressures

To successfully reproduce, many carnivorous mammals need access to suitable den sites. Den site selection is often based on fitness related criteria like escape from predators, food availability and shelter from extreme weather conditions. African wild dogs are cooperative breeders that use a den to give birth to their offspring. They often co-exist with lions and spotted hyenas, both of which are known to kill African wild dog pups. Little is known about den site selection by African wild dogs. In this study, we compared vegetation characteristics and distribution of roads and waterholes around den sites and random sites, in areas with high and low lion and spotted hyena densities. In both areas, African wild dogs selected den sites in closed woodland with little visibility, which is likely to reduce detection by predators, increase the likelihood of escape when detected, and might provide shelter from extreme weather conditions. In the high predator density area, African wild dogs seemed to spatially avoid predators by selecting den sites in this type of habitat relatively further away from waterholes and roads. African wild dogs have high energetic costs of gestation. Therefore, even when predation risk is relatively low, they are likely to try to maximise their fitness by choosing a den site in habitat that will provide optimal protection for their offspring, leaving little additional options to respond to a higher predation pressure.     

Foraging costs,hunting success and its implications for African wild dog (Lycaon pictus) conservation inside and outside a protected area

When selecting a habitat, animals utilize habitat in which they yield the highest rate of energy. Differences in foraging costs and hunting success are therefore likely to affect habitat choice. In a previous study, we showed that African wild dog (Lycaon pictus) packs with territories inside Hwange National Park (HNP), over the course of several years, moved their territories into the buffer zone outside HNP, where reproductive success was higher but anthropogenic mortality exceeded natality. In this study, based on long‐term radio‐telemetry data from 22 African wild dog packs, we analysed whether differences in foraging costs and hunting success could have contributed to this territorial drift. Taking seasonality and pack size into account, we determined foraging costs (foraging distance and chase distance) and hunting success (successful or failed chase) inside and outside HNP. Although we observed no difference in foraging costs, hunting success was higher outside HNP, which is likely to have contributed to the territorial drift into the buffer zone outside the protected area. This study shows the importance of taking factors affecting hunting success into account in the conservation strategy of African wild dogs.     

African wild dogs test the 'survival of the fittest' paradigm

Charles Darwin first used the term 'survival of the fittest' in the 5th edition of The origin of species. A literal interpretation implies that predators will selectively prey upon the weakest members of a population. We demonstrate that this is true for African wild dogs hunting impala.     

Dynamics of a small re-introduced population of wild dogs over 25 years: Allee effects and the implications of sociality for endangered species’ recovery

We analysed 25 years (1980–2004) of demographic data on a small re-introduced population of endangered African wild dogs (Lycaon pictus) in Hluhluwe-iMfolozi Park (HiP), South Africa, to describe population and pack dynamics. As small populations of cooperative breeders may be particularly prone to Allee effects, this extensive data set was used to test the prediction that, if Allee effects occur, aspects of reproductive success, individual survival and population growth should increase with pack and population size. The results suggest that behavioural aspects of wild dogs rather than ecological factors (i.e. competitors, prey and rainfall) primarily have been limiting the HiP wild dog population, particularly a low probability of finding suitable mates upon dispersal at low pack number (i.e. a mate-finding Allee effect). Wild dogs in HiP were not subject to component Allee effects at the pack level, most likely due to low interspecific competition and high prey availability. This suggests that aspects of the environment can mediate the strength of Allee effects. There was also no demographic Allee effect in the HiP wild dog population, as the population growth rate was significantly negatively related to population size, despite no apparent ecological resource limitation. Such negative density dependence at low numbers indicates that behavioural studies of the causal mechanisms potentially generating Allee effects in small populations can provide a key to understanding their dynamics. This study demonstrates how aspects of a species’ social behaviour can influence the vulnerability of small populations to extinction and illustrates the profound implications of sociality for endangered species’ recovery. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial and temporal avoidance of risk within a large carnivore guild

Within a large carnivore guild, subordinate competitors (African wild dog, Lycaon pictus, and cheetah, Acinonyx jubatus) might reduce the limiting effects of dominant competitors (lion, Panthera leo, and spotted hyena, Crocuta crocuta) by avoiding them in space, in time, or through patterns of prey selection. Understanding how these competitors cope with one other can inform strategies for their conservation. We tested how mechanisms of niche partitioning promote coexistence by quantifying patterns of prey selection and the use of space and time by all members of the large carnivore guild within Liuwa Plain National Park in western Zambia. Lions and hyenas specialized on wildebeest, whereas wild dogs and cheetahs selected broader diets including smaller and less abundant prey. Spatially, cheetahs showed no detectable avoidance of areas heavily used by dominant competitors, but wild dogs avoided areas heavily used by lions. Temporally, the proportion of kills by lions and hyenas did not detectably differ across four time periods (day, crepuscular, early night, and late night), but wild dogs and especially cheetahs concentrated on time windows that avoided nighttime hunting by lions and hyenas. Our results provide new insight into the conditions under which partitioning may not allow for coexistence for one subordinate species, the African wild dog, while it does for cheetah. Because of differences in responses to dominant competitors, African wild dogs may be more prone to competitive exclusion (local extirpation), particularly in open, uniform ecosystems with simple (often wildebeest dominated) prey communities, where spatial avoidance is difficult.     

Dogs Canis familiaris as carnivores: their role and function in intraguild competition

• 1 Dogs Canis familiaris are the world's most common carnivore and are known to interact with wildlife as predators, prey, competitors, and disease reservoirs or vectors.
• 2 Despite these varied roles in the community, the interaction of dogs with sympatric wild carnivore species is poorly understood. We review how dogs have been classified in the literature, and illustrate how the location and ranging behaviour of dogs are important factors in predicting their interactions with wild prey and carnivores.
• 3 We detail evidence of dogs as intraguild competitors with sympatric carnivores in the context of exploitative, interference and apparent competition.
• 4 Dogs can have localized impacts on prey populations, but in general they are not exploitative competitors with carnivores. Rather, most dog populations are highly dependent on human‐derived food and gain a relatively small proportion of their diet from wild prey. However, because of human‐derived food subsidies, dogs can occur at high population densities and thus could potentially outcompete native carnivores, especially when prey is limited.
• 5 Dogs can be effective interference competitors, especially with medium‐sized and small carnivores. Dogs may fill the role of an interactive medium‐sized canid within the carnivore community, especially in areas where the native large carnivore community is depauperate.
• 6 Dogs can also be reservoirs of pathogens, because most populations around the world are free‐ranging and unvaccinated. Diseases such as rabies and canine distemper have resulted in severe population declines in several endangered carnivores coexisting with high‐density dog populations. Dogs can therefore be viewed as pathogen‐mediated apparent competitors, capable of facilitating large‐scale population declines in carnivores.
• 7 Based on this information, we propose conceptual models that use dog population size and ranging patterns to predict the potential for dogs to be intraguild competitors. We discuss how interactions between dogs and carnivores might influence native carnivore communities.

     

Inferring the ancestry of African wild dogs that returned to the Serengeti-Mara

An endangered population of African wild dogs (Lycaon pictus) disappeared from the Serengeti-Mara area in 1991. The reasons for the extinction are not well understood, but disease was implicated in the disappearance. In 2001, wild dogs naturally re-established themselves in the region. We conducted genetic profiling on samples collected prior and subsequent to this event, as well as samples from three geographically close populations, to determine the potential source of colonisers. Contrary to expectations, we found no evidence of re-colonisation from these nearby wild dog populations. Rather, our analyses suggest that the re-established animals are primarily derived from the same genetic population as the pre-extinction animals, indicating that wild dogs are likely to have persisted in the Serengeti-Mara after 1991. We also detected some migrants that could be derived from genetically distinct populations outside the recovery area. Overall, we did not detect a decline in genetic diversity at either neutral microsatellites or major histocompatibility complex loci, indicating that the supposed disappearance of wild dogs in the Serengeti-Mara did not substantially impact genetic variation of the population.     

Evaluating the Status of and African Wild Dogs Lycaon pictus and Cheetahs Acinonyx jubatus through Tourist-based Photographic Surveys in the Kruger National Park

The Kruger National Park is a stronghold for African wild dog Lycaon pictus and cheetah Acinonyx jubatus conservation in South Africa. Tourist photographic surveys have been used to evaluate the minimum number of wild dogs and cheetahs alive over the last two decades. Photographic-based capture-recapture techniques for open populations were used on data collected during a survey done in 2008/9. Models were run for the park as a whole and per region (northern, central, southern). A total of 412 (329–495; SE 41.95) cheetahs and 151 (144–157; SE 3.21) wild dogs occur in the Kruger National Park. Cheetah capture probabilities were affected by time (number of entries) and sex, whereas wild dog capture probabilities were affected by the region of the park. When plotting the number of new individuals identified against the number of entries received, the addition of new wild dogs to the survey reached an asymptote at 210 entries, but cheetahs did not reach an asymptote. The cheetah population of Kruger appears to be acceptable, while the wild dog population size and density are of concern. The effectiveness of tourist-based surveys for estimating population sizes through capture-recapture analyses is shown.     

Diet choice of large carnivores in northern Cameroon

The diet of African wild dogs, lions and spotted hyenas was studied in the woodland savannah of Faro National Park and surrounding hunting zones in northern Cameroon during the dry season. Faecal analysis revealed that the diet of the three large carnivores overlaps considerably. Frequencies of prey remains of large and medium‐sized antelopes were highest, with Buffon's kob being by far the most common prey species. Carnivores seem to consume most abundant prey species. No prey item of domestic livestock was found in the faeces. Faecal analysis has proved to be a useful method to reveal diet choice for large carnivores in northern Cameroon. Detailed information on the distribution and density of large carnivores and their prey is needed to give a better picture of the status of carnivores in northern Cameroon and to help resolve conflicts with livestock. Further investigations into the seasonal variation of predation and other factors structuring the large carnivore community in northern Cameroon are recommended.     

Diet choice of large carnivores in northern Cameroon

The diet of African wild dogs, lions and spotted hyenas was studied in the woodland savannah of Faro National Park and surrounding hunting zones in northern Cameroon during the dry season. Faecal analysis revealed that the diet of the three large carnivores overlaps considerably. Frequencies of prey remains of large and medium‐sized antelopes were highest, with Buffon's kob being by far the most common prey species. Carnivores seem to consume the most abundant prey species. No prey item of domestic livestock was found in the faeces. Faecal analysis has proved to be a useful method to reveal diet choice for large carnivores in northern Cameroon. Detailed information on the distribution and density of large carnivores and their prey is needed to give a better picture of the status of carnivores in northern Cameroon and to help resolve conflicts with livestock. Further investigations into the seasonal variation of predation and other factors structuring the large carnivore community in northern Cameroon are recommended.     

African Wild Dog Dispersal and Implications for Management

Successful conservation of species that roam and disperse over large areas requires detailed understanding of their movement patterns and connectivity between subpopulations. But empirical information on movement, space use, and connectivity is lacking for many species, and data acquisition is often hindered when study animals cross international borders. The African wild dog (Lycaon pictus) exemplifies such species that require vast undisturbed areas to support viable, self-sustaining populations. To study wild dog dispersal and investigate potential barriers to movements and causes of mortality during dispersal, between 2016 and 2019 we followed the fate of 16 dispersing coalitions (i.e., same-sex group of ≥1 dispersing African wild dogs) in northern Botswana through global positioning system (GPS)-satellite telemetry. Dispersing wild dogs covered ≤54 km in 24 hours and traveled 150 km to Namibia and 360 km to Zimbabwe within 10 days. Wild dogs were little hindered in their movements by natural landscape features, whereas medium to densely human-populated landscapes represented obstacles to dispersal. Human-caused mortality was responsible for >90% of the recorded deaths. Our results suggest that a holistic approach to the management and conservation of highly mobile species is necessary to develop effective research and evidence-based conservation programs across transfrontier protected areas, including the need for coordinated research efforts through collaboration between national and international conservation authorities. © 2020 The Wildlife Society.     

Rabies Virus and Canine Distemper Virus in Wild and Domestic Carnivores in Northern Kenya: Are Domestic Dogs the Reservoir?

Rabies virus (RV) and canine distemper virus (CDV) can cause significant mortality in wild carnivore populations, and RV threatens human lives. We investigated serological patterns of exposure to CDV and RV in domestic dogs (Canis familiaris), African wild dogs (Lycaon pictus), black-backed jackals (Canis mesomelas), spotted hyenas (Crocuta crocuta), striped hyenas (Hyaena hyaena) and African lions (Panthera leo), over a 10-year period, in a Kenyan rangeland to assess the role domestic dogs may play in the transmission dynamics of these two important canid pathogens. Observed patterns of RV exposure suggested that repeated introduction, rather than maintenance, occurred in the wild carnivore species studied. However, RV appeared to have been maintained in domestic dogs: exposure was more likely in domestic dogs than in the wild carnivores; was detected consistently over time without variation among years; and was detected in juveniles (≤1-year-old) as well as adults (>1-year-old). We conclude that this domestic dog population could be a RV reservoir. By contrast, the absence of evidence of CDV exposure for each carnivore species examined in the study area, for specific years, suggested repeated introduction, rather than maintenance, and that CDV may require a larger reservoir population than RV. This reservoir could be a larger domestic dog population; another wildlife species; or a “metareservoir” consisting of multiple interconnected carnivore populations. Our findings suggest that RV risks to people and wild carnivores might be controlled by domestic dog vaccination, but that CDV control, if required, would need to target the species of concern.     

African Wild Dogs as a Fugitive Species: Playback Experiments Investigate How Wild Dogs Respond to their Major Competitors

It has been suggested that African wild dogs Lycaon pictus need exceptionally large home ranges (and hence occur at such low densities) because they are limited by competition with larger sympatric carnivores, namely lions Panthera leo and spotted hyenas Crocuta crocuta. To investigate this relationship at a proximate level and explore which factors mediate it, we conducted audio playback experiments examining how wild dogs responded to the simulated proximity of either lions or hyenas. The principle finding was that wild dogs consistently moved directly away from lion roars, but when played hyena whoops either stood their ground or, later, moved off in a random direction. These results suggest that lions represent an immediate high‐level threat to wild dogs that is invariably best avoided, whilst the threat from hyenas may not be so great or perhaps is simply unavoidable. Wild dogs appeared to make some assessment of ambush risk during interactions with lions, illustrated by the varying latency to their retreat in habitats of differing vegetation density (and hence ambush potential). Additionally, packs with younger pups were more likely to alarm call and exhibited a slower rate of retreat in the hour following exposure to lion roars. Other variables investigated (competitor group size, lion sex, presence of pups) failed to explain variation in wild dogs’ responses.     

The effects of anthrax on endangered African wild dogs (Lycaon pictus)

African wild dogs ( Lycaon pictus ) are large, social canids formerly found throughout sub-Saharan Africa. Wild dogs are now endangered; only six nations hold populations larger than 100 individuals. Considerable evidence suggests that disease plays a central role in the regulation of some wild dog populations. In particular, it has been suggested that epizootics of rabies, canine distemper and anthrax can have strong local effects on wild dog numbers. Resolution of the regulatory importance of these diseases has been hampered by lack of data from wild populations. Here we report on an outbreak of anthrax among wild dogs in the Selous Game Reserve in southern Tanzania, describing clinical signs and diagnosis, measuring mortality, and testing for effects on hunting success and movements. Only one of five packs under study was affected. Immediate mortality was mild, with none of 18 adults (0%) and four of 24 pups (17%) dying. Mortality was significantly higher among puppies (<1 year old) than among adults. Mortality of individuals that showed signs of disease but did not die immediately was not significantly elevated over the following six months. The hunting success and movement patterns of the pack were not affected by the outbreak. Collectively, these data suggest that African wild dogs possess a degree of resistance to anthrax.     

Seasonal regulation of the growth hormone-insulin-like growth factor-I axis in the American black bear (Ursus americanus)

The American black bear maintains lean body mass for months without food during winter denning. We asked whether changes in the growth hormone/insulin-like growth factor-I (GH-IGF-I) axis may contribute to this remarkable adaptation to starvation. Serum IGF-I levels were measured by radioimmunoassay, and IGF-binding proteins (IGFBPs) were analyzed by ligand blotting. Initial studies in bears living in the wild showed that IGF-I levels are highest in summer and lowest in early winter denning. Detailed studies in captive bears showed that IGF-I levels decline in autumn when bears are hyperphagic, continue to decline in early denning, and later rise above predenning levels despite continued starvation in the den. IGFBP-2 increased and IGFBP-3 decreased in early denning, and these changes were also reversed in later denning. Treatment with GH (0.1 mg·kg(-1)·day(-1) × 6 days) during early denning increased serum levels of IGF-I and IGFBP-3 and lowered levels of IGFBP-2, indicating that denning bears remain responsive to GH. GH treatment lowered blood urea nitrogen levels, reflecting effects on protein metabolism. GH also accelerated weight loss and markedly increased serum levels of free fatty acids and β-hydroxybutyrate, resulting in a ketoacidosis (bicarbonate decreased to 15 meq/l), which was reversed when GH was withdrawn. These results demonstrate seasonal regulation of GH/IGF-I axis activity in black bears. Diminished GH activity may promote fat storage in autumn in preparation for denning and prevent excessive mobilization and premature exhaustion of fat stores in early denning, whereas restoration of GH/IGF activity in later denning may prepare the bear for normal activity outside the den.     

Movement and Habitat Use of Canada Lynx During Denning in Minnesota

Abstract Establishing whether conditions are suitable for reproduction would help determine if immigration is necessary for Canada lynx (Lynx canadensis) to persist at the southern edge of the species range. We located den sites and monitored reproduction of radiocollared lynx in Minnesota from 2004 to 2006. Movement rates of denning females measured with Global Positioning System collars were similar to movement rates of lynx elsewhere. Female lynx with kittens used different habitat types in predenning, denning, and postdenning periods. Landscape composition at the scale of the foraging radius around a den site contained more lowland conifer, upland conifer, and regenerating forest than did home ranges or the area used by radiocollared lynx in Minnesota, USA. We used the spatial distribution of cover-type composition around known den sites to predict and map potential denning habitat in northeastern Minnesota. Techniques for identifying the spatial distribution of suitable denning habitat provide a biological basis for management actions that could enhance recovery of Canada lynx populations in the southern part of the species range.     

Changing vulnerability to predation related to season and sex in an African ungulate assemblage

The consequences of predation for prey population dynamics depend on the extent to which this mortality is predisposed by malnutrition or senescence, or additive in the sense that animals that would otherwise not have died at that time were killed. In places lacking effective predators, few adult ungulates die during the summer or wet season months when food is plentifully available. Hence the seasonal distribution of predator kills as well as the age and sex classes of the prey indicates the extent to which malnutrition contributes to mortality as well as other influences on vulnerability. Using records of animal deaths assembled over 35 years in South Africa's Kruger National Park, these patterns were investigated for 12 ungulate species forming the prey of lions, and for three other large predators with respect to one prey species. Buffalo, kudu and giraffe were more strongly represented in kills made during the late dry season, while wildebeest and zebra made relatively greater contributions during the wet season. Impala, waterbuck, warthog and rarer antelope species became more prominent in kills during transitional periods between seasons. Five prey species showed an elevation in representation of males in lion kills during the mating season, as well as impala for all predator species. Females were more prominently represented in kills during the time of late gestation and parturition for three prey species. Hence reproductive activities as well as changing vegetation cover and food resources affected vulnerability to predation. Shifts in susceptibility to predation over the seasonal cycle corresponded with rainfall‐related variation in the annual representation of these ungulate species in lion kills. The availability of vulnerable prey species, age and sex classes at different stages of the seasonal cycle helps maintain a high abundance of lions. These factors contribute to the strong additive impact that predation has had on the abundance of some of these ungulate populations.