Landowners’ perspectives of black‐backed jackals (Canis mesomelas) on farmlands in KwaZulu‐Natal,South Africa

Despite continued efforts to eradicate black‐backed jackals (Canis mesomelas), they are considered an abundant mesopredator on agricultural land across South Africa, resulting in ongoing human–wildlife conflict and concern for farmers and wildlife managers. We conducted a questionnaire survey and semi‐formal interviews with farmers throughout KwaZulu‐Natal, examining farmers’ livestock husbandry, land‐use changes and perspectives towards jackals as a perceived threat to livestock. Many (75%) respondents acknowledged expanding agricultural activities on their farmlands since the onset of their farming careers. However, the perception was that these changes placed little pressure on mesopredators as farmers reported frequent daily (25%) and weekly (31%) sightings of jackal, and regular predation on livestock (72%). Some landowners (31%) reported between one and five livestock losses annually and suggest that mitigation strategies to prevent livestock losses are in place. Farmers suggested the increasing intensity in agricultural practices provided a greater food source for jackals allowing them to thrive in expanding agricultural conditions and, in some circumstances, farmers admitted to possibly being a cause through poor disposal techniques for dead animals. Feedback from farmers emphasized the importance of having collaboration between farmers to control jackal predation and reduce human–wildlife conflict.     

Diet,Prey Selection,and Predation Impact of Black-Backed Jackals in South Africa

ABSTRACT To investigate the role of black-backed jackals (Canis mesomelas) as predators, we studied diet, prey selection, and predation impact of jackals on 2 game ranches in South Africa that differed in ungulate diversity and biomass. Results showed that large (>15 kg) ungulate species dominated jackal diets throughout the year on both the less diverse (range of ingested biomass across seasons = 39–78%) and more diverse (26–69%) game ranch. Other important food items included medium-sized mammals (1–3 kg; 1–26%) and fruit (2–69%), whereas small mammals comprised 3–11% of ingested biomass across seasons on both sites. Jackals were not random in consumption of ungulates, and consumption patterns suggested jackals actively hunted certain species rather than consumed them as carrion. During ungulate birthing periods, jackals consumed almost exclusively those ungulate species that were hiders (i.e., fawns were hidden in tall vegetation away from herd) regardless of ungulate densities, suggesting that primarily fawns were preyed upon. Among hiders, there was a negative relationship (P = 0.01) between body size and percent of population consumed by jackals, indicating smaller species were more susceptible than larger species to jackal predation. Consequently, springbok (Antidorcas marsupialis) were always selected over other ungulate species on both sites, and this species was the most impacted by jackal predation. In contrast, ungulate species that were followers (i.e., fawns immediately followed mothers within protection of the herd) were scarcely or not at all consumed by jackals, regardless of body size or density. Medium-sized mammals were selectively consumed over ungulates, and there was a negative relationship (P < 0.01) between consumption of berries and ungulates, indicating alternative food resources influenced consumption of ungulates on our study sites. Our results will help wildlife managers in Africa identify ungulate species susceptible to jackal predation, and can be used to develop management strategies for reducing jackal predation in areas where it is problematic.     

Social organization,home ranges,and extraterritorial forays of black-backed jackals

We radio-tracked 15 black-backed jackals (Canis mesomelas) from 8 adjacent family groups on Benfontein Game Farm (i.e., Benfontein) in South Africa to investigate their movement patterns and social organization. Jackal family groups consisted of mated pairs (alphas), 0–3 nonbreeding adults (betas), and pups, depending on the season. Mean (±SE) home-range size of alphas (9.4 ± 1.2 km², n = 6) did not differ (P = 0.766) from betas (9.8 ± 0.7 km², n = 8). Most beta jackals (8 of 10) remained philopatric on Benfontein, apparently because of the high density of springbok (Antidorcas marsupialis), their preferred prey. Three of 5 alphas and all 8 betas went on extraterritorial forays (i.e., forays). Generally, betas spent more of their active time on forays (2–20% of time) than alphas (0–3%; P = 0.048), and betas went farther on forays (2–8 km) than alphas (2–3 km; P = 0.003). The number of forays differed (P < 0.001) among seasons; most forays occurred during summer (64%) when jackals visited neighboring livestock farms, apparently to predate on domestic sheep. Overall, our results indicate forays by jackals are affected by social status, seasonal availability of preferred prey, and the reproductive cycle of jackals. To reduce jackal predation on livestock farms near reserves, we recommend that preventative measures (e.g., use of herders, jackal control activities) be increased during summer when jackals are most likely to travel outside reserves. © 2019 The Wildlife Society.     

Resource partitioning among cape foxes,bat-eared foxes,and black-backed jackals in South Africa

Cape foxes (Vulpes chama) and bat-eared foxes (Otocyon megalotis) are sympatric with black-backed jackals (Canis mesomelas) over much of southern Africa, although competition with and/or predation by jackals may suppress local populations of both fox species. From 2005 to 2008, we captured, radio-collared, and monitored 11 cape foxes, 22 bat-eared foxes, and 15 black-backed jackals on a game ranch in South Africa to investigate their spatial, habitat, temporal, and dietary resource overlap. Mean annual home-range sizes were 27.7 km² for cape foxes, 5.0 km² for bat-eared foxes, and 17.8 km² for jackal family groups. Home ranges overlapped completely between species, although core areas overlapped less (<45%), with cape foxes and jackals overlapping the least (12%). When active, cape foxes, but not bat-eared foxes, used core areas of jackal groups less than expected. Additionally, both fox species used jackal core areas less than expected for their den sites, suggesting areas outside jackal core areas were used as refuges by foxes. Strong levels of habitat partitioning were not apparent at the study site or home-range levels, although habitat selection for den sites differed between jackals and cape foxes. Jackals were the most diurnal across seasons, whereas cape foxes were the most nocturnal. Diets overlapped little (R₀ = 0.20–0.34) among the canid species, with bat-eared foxes overlapping the least with the others. Jackals killed at least 5 collared bat-eared foxes and 1 collared cape fox, indicating potential interference competition, probably for exclusive use of territorial space rather than over shared resources. We conclude that bat-eared foxes coexisted with jackals primarily by their dietary specialization and group living. Cape foxes coexisted with jackals by exhibiting high levels of spatial, habitat, temporal, and dietary partitioning. Surprisingly, the fox species exhibited positive associations with each other. Our results show the mechanisms that may allow jackals to suppress fox populations, yet also show how foxes, in turn, use different mechanisms to coexist with a dominant canid. © 2012 The Wildlife Society.     

Black-backed jackal diet at Mokolodi Nature Reserve, Botswana

We examined the feeding habits of black‐backed jackals at Mokolodi Nature Reserve, Botswana, by analysing 237 scats collected between November 1995 and February 1997. Jackal dietary habits reflected the availability of a wide variety of food items and the differential vulnerability of prey. Potential animal and plant food available to jackals varies throughout the year because of its seasonal character. Seasonality of prey occurrence in scats was pronounced for small mammals, miscellaneous fruits and invertebrates. Across all seasons, mammals were the most common food resource (32.4%, n = 168), followed by anthropogenic items (14.8%), fruits (12.9%), invertebrates (10.8%), birds (8.5%), unidentified items (3.5%) and reptiles (1.4%). The presence of domestic mammals and poultry remains in scats reveals their importance in the diet of jackals and the tendency of jackals to frequent human settlements in search of food. Some ecological implications of jackal dietary habits are also explored.     

Seasonal diet and prey selection of black‐backed jackals on a small‐livestock farm in South Africa

The extent to which black‐backed jackals (Canis mesomelas) selectively consume domestic sheep (Ovis aries) compared to wild prey is unknown. Using faecal analysis and prey surveys, we determined the seasonal diet and prey selection of jackals on a small‐livestock farm in South Africa. Sheep comprised 25–48% of the biomass consumed by jackals across seasons, and consumption peaked during the lambing seasons, indicating sheep often were the main food resource for jackals. Another main food resource was wild ungulates <50 kg, primarily springbok (Antidorcas marsupialis) and steenbok (Raphicerus campestris), which comprised 8–47% of the biomass consumed. Other important food items were mammals 1–3 kg (4–16%), which included hares (Lepus spp.) and springhares (Pedetes capensis), and small rodents (10–14%). Compared to the biomass available, jackals selectively consumed mammals 1–3 kg over sheep across all seasons, whereas wild ungulates <50 kg were selectively consumed over sheep in most seasons. Our results showed that jackals selectively consumed different food items throughout the year and that wild prey were consistently selected over sheep.     

Rabies in Zimbabwe: reservoir dogs and the implications for disease control.

Using detailed field study observations of the side-striped jackal (Canis adustus) and a simple stochastic model of the transmission dynamics of the virus and host demography, we discuss the epidemiology of rabies virus infection in the jackal population of Zimbabwe. Of the two jackal species in Zimbabwe, the other being the black-backed jackal (Canis mesomelas), the bulk of notified rabies cases are in side-striped jackals. Specifically, we show that the side-striped jackal population itself does not seem able to support rabies infection endemically, i.e. without frequent reintroduction from outside sources of infection. We argue that this is probably because the overall average jackal population density is too low to maintain the chain of infection. This study suggests that the disease is regularly introduced to jackals by rabid dogs from populations associated with human settlements. Given the rapidly rising dog population in Zimbabwe, estimates are derived of the future incidence of jackal rabies based on different dog-vaccination scenarios.     

Short‐term foraging responses of a generalist predator to management‐driven resource pulses

Generalist predators characteristically switch prey in response to resource pulses. Ungulate population reinforcement through translocation provides stressed and vulnerable prey, and apex predator reintroduction provides carrion, both resources that can be exploited by generalist predators. We investigated the dietary response of black‐backed jackal to two management interventions in the Karoo National Park, South Africa, namely the reinforcement of the springbok population, and then the reintroduction of lions. By analysing jackal diets from scats collected before and after each management intervention, we show that the relative per cent occurrence and biomass consumed of springbok increased in the diet following the population reinforcement of springbok. In contrast, large ungulates were more prevalent in the diet following apex predator reintroduction. These results suggest that jackals took advantage of a potentially vulnerable abundant springbok population following their population reinforcement, and then switched to foraging on carrion provided by reintroduced lions. These results provide insights into the dietary response of a generalist mesopredator to management interventions. A key lesson from the study is that the diet of generalist predators is context‐dependent and should be interpreted in that light.     

A European Concern? Genetic Structure and Expansion of Golden Jackals (Canis aureus) in Europe and the Caucasus

In the first continent-wide study of the golden jackal (Canis aureus), we characterised its population genetic structure and attempted to identify the origin of European populations. This provided a unique insight into genetic characteristics of a native carnivore population with rapid large-scale expansion. We analysed 15 microsatellite markers and a 406 base-pair fragment of the mitochondrial control region. Bayesian-based and principal components methods were applied to evaluate whether the geographical grouping of samples corresponded with genetic groups. Our analysis revealed low levels of genetic diversity, reflecting the unique history of the golden jackal among Europe’s native carnivores. The results suggest ongoing gene flow between south-eastern Europe and the Caucasus, with both contributing to the Baltic population, which appeared only recently. The population from the Peloponnese Peninsula in southern Greece forms a common genetic cluster with samples from south-eastern Europe (ΔK approach in STRUCTURE, Principal Components Analysis [PCA]), although the results based on BAPS and the estimated likelihood in STRUCTURE indicate that Peloponnesian jackals may represent a distinct population. Moreover, analyses of population structure also suggest either genetic distinctiveness of the island population from Samos near the coast of Asia Minor (BAPS, most STRUCTURE, PCA), or possibly its connection with the Caucasus population (one analysis in STRUCTURE). We speculate from our results that ancient Mediterranean jackal populations have persisted to the present day, and have merged with jackals colonising from Asia. These data also suggest that new populations of the golden jackal may be founded by long-distance dispersal, and thus should not be treated as an invasive alien species, i.e. an organism that is “non-native to an ecosystem, and which may cause economic or environmental harm or adversely affect human health”. These insights into the genetic structure and ancestry of Baltic jackals have important implications for management and conservation of jackals in Europe. The golden jackal is listed as an Annex V species in the EU Habitats Directive and as such, considering also the results presented here, should be legally protected in all EU member states.     

The golden jackal (<Emphasis Type="Italic">Canis aureus</Emphasis>) in Bosnia and Herzegovina: density of territorial groups,population trend and distribution range

In this article, we present the results of the first systematic surveys of golden jackals in Bosnia and Herzegovina (B&H). The population status and distribution of the golden jackal (Canis aureus) in B&H was largely unknown, and the few existing literature records mention their presence at only five localities in the country. We interviewed managers of all the hunting grounds in B&H and reviewed available jackal hunting records from 2000 to 2016. In total, we collected 212 records of legally harvested jackals. We observed an increasing trend of harvested jackals in B&H (on average 35% annual increase) during this same period. Using the acoustic (playback) method, we confirmed the presence of 80 territorial jackal groups along six transects covering 3081 km². We estimated density to be a minimum of 0.33 groups/10 km² in northern B&H and 0.10 groups/10 km² in central B&H. Jackal groups were located at significantly lower altitudes with respect to available area along the transects. We present a distribution map of confirmed jackal occurrences in B&H, which indicates that the core area of jackal distribution in the country is currently located along the Sava River and its tributaries in the northern part of B&H. Jackals are sporadically present in the rest of the country, where gray wolves (Canis lupus) probably limit their presence. In total, jackal presence has been detected in 19% (109 out of 586) of 10?×?10 km grid cells covering the country. The primary factor driving the expanding population of jackals in northern B&H appears to be immigration of jackals from Croatia and Serbia.     

Population densities and habitat use of the golden jackal (Canis aureus) in farmlands across the Balkan Peninsula

The main objective of this study was to analyze the habitat use and population densities of the golden jackal in four countries across lowland regions of the Balkan Peninsula, known as the core area of the species' distribution in Europe. Using indirect (acoustic) method for detecting territorial golden jackals, we surveyed jackal presence and densities on 331 monitoring sites in four countries, covering area an of 4,296 km² in total during April and May 2007–2012. We used GIS to assess landscape and environmental characteristics in a 2-km circular buffer (12.6 km²) around calling stations. Average population density of golden jackals in the study areas ranged between 0.6 and 1.1 territorial groups/10 km² (mean ± SE, 0.6?±?0.06 groups/10 km²), with several high-density areas with up to 4.8 territorial groups/10 km². Analysis of habitat use showed that for both jackal occurrence and number of jackal groups, the only significant parameter was the interaction between country and intensity of agriculture, indicating that jackals adapt their habitat selection patterns in relation to the habitat availability. We observed that selection of the more suitable habitats (shrub–herbaceous vegetation/heterogeneous agricultural vegetation) increased with lower proportion of these habitat types in the study area. Our study confirms high habitat plasticity of the golden jackal and offers explanation for its recent range expansion, which might be connected with the land use changes during the last decades in the Balkan Peninsula.     

Genetic characterization of populations of the golden jackal and the red fox in Israel

The golden jackal and red fox are among the wildlife species protected by Israeli law as enforced by the Israel Nature and Parks Authority. In 1964, as a part of a management program to control rabies in Israel, a poison eradication campaign was launched to exterminate golden jackals, considered to be the main reservoir of the disease. The program resulted in the near-complete extermination of jackals in Israel, while foxes were only mildly affected. Jackals have since regained their original numbers and have recolonized southern Israel. We here examined the population structure of the golden jackal and red fox in Israel, 48 years after the poison eradication campaign. DNA from 88 golden jackals and 89 red foxes representing five different geographic regions was extracted and amplified at 13 microsatellite loci in order to characterize the populations on a genetic level. High genetic diversity was found among the jackal and fox populations. A possible migration route through the Jordan Rift Valley was suggested for both species by the genetic similarity of populations in northern and southern Israel. However, in both species, the animals from the center of Israel were distinctive from those north or south, indicating the relative isolation of central populations, likely due to fragmentation or a high abundance of food resources. Genetic profiles obtained for the golden jackal and the red fox in Israel may aid in their conservation management and in the study of zoonotic diseases.     

Spatial ecology of golden jackal in farmland in the Ethiopian Highlands

The spatial ecology of golden jackal Canis aureus was studied on farmland adjacent to the Bale Mountains National Park in southern Ethiopia during 1998–2000. Three adult and four subadult jackals were captured in leg‐hold traps and radiotagged. The range size of the adult jackals varied from 7.9 to 48.2 km² and the subadults from 24.2 to 64.8 km² . These ranges are the largest recorded for this species. Range overlap of the tagged jackals averaged 54%, which, in conjunction with observations of associations between individuals, suggested that all the tagged jackals belonged to one social group. Tagged jackals were observed alone on 87% of occasions despite the extensive overlap in individual ranges. Pairs consisting of a male and female were the most commonly observed group and larger groups were seen on only five occasions. Jackals in this population appeared less gregarious than observed elsewhere. The jackals used all the habitats available to them, particularly at night when they foraged in Artemesia and Hypericum bush and farmland. During the day they were more frequently found in Hagenia and Juniper woodland and their diurnal resting sites were characterized by thick cover. This is the first detailed study of golden jackals in a human‐modified landscape in Africa and further demonstrates the flexibility in behaviour and ecology exhibited by this species throughout its range.     

Genetic structure and expansion of golden jackals (Canis aureus) in the north-western distribution range (Croatia and eastern Italian Alps)

The golden jackal, widely distributed in Europe, Asia and Africa, is one of the less studied carnivores in the world and the genetic structure of the European populations is unknown. In the last century jackals strongly declined mainly due to human persecution, but recently they expanded again in eastern Europe. With the aim to determine the genetic structure and the origin of expanding jackals, we analyzed population samples obtained from Bulgaria, Serbia, Croatia (Dalmatia and Slavonia) and individuals sampled in north-eastern Italy. Samples were typed at the hypervariable part of the mitochondrial DNA control-region (mtDNA CR1) and at 15 canine autosomal microsatellite loci (STR), and analyzed using multivariate, Bayesian and landscape genetic methods. The mtDNA CR1 was monomorphic, showing a single haplotype shared among all the populations. The STR loci were variable, with 2–14 alleles and intermediate values of heterozygosity (Ho = 0.47; He = 0.51). Genetic diversity was significantly partitioned (θ_ST = 0.07; P < 0.001) and the populations were partially distinct, perhaps in consequence of recent fragmentations. Jackals from Dalmatia were the most genetically differentiated. Assignment testing and gene flow analyses suggested that jackals colonizing Italy have admixed origins from Dalmatian and Slavonian populations. They are not first generation migrants, suggesting that dispersal towards north-eastern Italy is a stepping-stone process. Golden jackal and wolf colonization patterns might be different, with prevalent short-distance dispersal in jackals versus prevalent long distance dispersal in wolves. The admixed origin of jackals in the Alps ensures abundant genetic variability, which may enhance adaptive fitness and expectancy of population growth. The intersections between Dinaric–Balkan and Eastern Alps are areas of population expansion and admixture, highlighting their conservation, ecological and evolutionary values.     

Late autumn trophic flexibility of the golden jackalCanis aureus

The feeding habits of the golden jackal Canis aureus (Linnaeus, 1758) were compared using scat analysis in Hungary (temperate climate agricultural area), Greece (Mediterranean marshland), and Israel (Mediterranean agricultural area). Samples (84, 70 and 64 scats, respectively) were collected during late autumn, a period with capital importance to the long term survival of young jackals, during which they become independent. Predation of wild-living prey species was highest in Hungary, consisting primarily of small mammals (biomass estimation: 51.5%, mainly rodents), contrary to Israel and Greece, where scavenging on domestic animals dominated the diet of jackals (74%, mainly poultry and 62.6%, mainly goats, respectively). The highest consumption of wild ungulates (mainly wild boar) was found in Greece (15.7%), and plants in Hungary (39%). Bird consumption was low in all three areas. Reptiles, amphibians and fish occurred only in the diet of jackals in Greece and Israel, whereas invertebrates were eaten more frequently in Hungary. Jackal dietary composition was extremely variable between regions, strongly associated with human presence. These results illustrate the golden jackal as having a very variable diet, resulting from opportunistic feeding habits.     

Phylogeography of the Golden Jackal (Canis aureus) in India

The golden jackal (Canis aureus) is one of the most common and widely distributed carnivores in India but phylogeographic studies on the species have been limited across its range. Recent studies have observed absence of mitochondrial (mt) DNA diversity in European populations while some North African populations of golden jackal were found to carry gray wolf (Canis lupus lupaster) mtDNA lineages. In the present study, we sequenced 440 basepairs (bp) of control region (CR) and 412 bp of cytochrome b (cyt b) gene of mtDNA from 62 golden jackals sampled from India (n = 55), Israel (n = 2) and Bulgaria (n = 5), to obtain a total of eighteen haplotypes, comprising sixteen from India and one each from Israel and Bulgaria. Except for three previously described haplotypes represented by one cyt b and one CR haplotype both from India, and one CR haplotype from Bulgaria, all haplotypes identified in this study are new. Genetic diversity was high in golden jackals compared to that reported for other canids in India. Unlike the paraphyletic status of African conspecifics with the gray wolf, the Indian (and other Eurasian) golden jackal clustered in a distinct but shallow monophyletic clade, displaying no evidence of admixture with sympatric and related gray wolf and domestic dog clades in the region. Phylogeographic analyses indicated no clear pattern of genetic structuring of the golden jackal haplotypes and the median joining network revealed a star-shaped polytomy indicative of recent expansion of the species from India. Indian haplotypes were observed to be interior and thus ancestral compared to haplotypes from Europe and Israel, which were peripheral and hence more derived. Molecular tests for demographic expansion confirmed a recent event of expansion of golden jackals in the Indian subcontinent, which can be traced back ~ 37,000 years ago during the late Pleistocene. Our results suggest that golden jackals have had a potentially longer evolutionary history in India than in other parts of the world, although further sampling from Africa, the Middle East and south-east Asia is needed to test this hypothesis.     

The ecology of large carnivores in the highlands of northern Ethiopia

The degradation and fragmentation of the northern Ethiopian highlands has resulted in frequent encounters of large carnivores with humans and their livestock. We interviewed 500 randomly selected households to estimate economic impact of livestock predation by spotted hyaena (Crocuta crocuta), leopard (Panthera pardus) and jackal (Canis aureus aureus) in the highlands of northern Ethiopia. The annual mean economic loss per household was approximately U.S.$ 20.2, about 7% of the average annual income of households in the area. Households surveyed reported losses of a total of 3122 livestock to hyaena, leopard and jackal predation over the past 5 years. This loss equated to a total financial loss of U.S.$ 50,381 . Livestock predation incidents of spotted hyaena, leopard and jackal demonstrated that spotted hyaena had a preference for dog, donkey, goat and sheep; leopard for goat, dog and sheep; and jackal for goat and sheep. Livestock predation of spotted hyaena and leopard were mainly during the night. We conclude that assessing depredation problems is important to develop actions for management of either livestock practices or wildlife conservation.     

Canine carnassials: character displacement in the wolves, jackals and foxes of Israel

Previous research implies that competitive character displacement in felids and mustelids of Israel is expressed by canine size. Anatomy and observed killing behaviour of canids suggest that canines in this group are less adapted for the stylized role they play in felids and mustelids. Thus we hypothesized that character displacement, if it exists in canids, should not be manifested more clearly by canine size than by other traits. Five sympatric and at least partially syntopic canids occupy Israel, while in North Africa the largest (wolf) and smallest (Blanford's fox) are absent. Sexual size dimorphism in Israeli canids is generally less than in felids and mustelids (in which we analysed each sex as a separate ‘morphospecies’), so we used mixed-sex samples to represent each species. The three largest species (wolf, golden jackal and red fox) are also represented by Middle Palaeolithic samples in Israel, and all three had larger carnassial lengths then. Carnassial lengths, canine diameters and skull lengths are all remarkable evenly spaced among the five recent species in Israel. In Egypt, no trait manifests significant equality. Despite regional fluctuations in size, the carnassial length ratios of the three smaller species (foxes) are strikingly constant (1.18–1.21) throughout the region, while the ratios for the three larger species (wolf, jackal and red fox), sympatric only in Israel, are larger (1.33–1.34). Finally, mean carnassial length of jackals is constant across North Africa, while skull length and canine diameter both increase from Algeria through Egypt. All three traits are larger in Egypt than in Israel. We tentatively ascribe the equal ratios in Israel to competitive character displacement, though this hypothesis is speculative because of numerous lacunae in knowledge of diet, killing behaviour, available resources and extent of food limitation. Furthermore, humans have greatly affected range, density and ecology of wolves and jackals in the last century. Larger sizes in the Palaeolithic may well be manifestations of Bergmann's rule. The constancy of carnassial length in North African jackals, notwithstanding a longitudinal cline in CBLs of these populations, and the constant ratio between jackal and red fox carnassial length are both consistent with a hypothesis of character release in the absence of the wolf.