Canis anthus

Geographic Range

African golden wolves ( Canis anthus ), which were considered the same species as Eurasian golden jackals ( Canis aureus ) until 2015, are found across northern Africa. Their range extends east to west from Somalia to Senegal and north to south from Algeria to Kenya. Thus, golden wolves occupy the Palearctic and Ethiopian faunal regions. Because golden wolves are a highly mobile species, their wide range was likely colonized naturally. Their historic range is unknown.

• Biogeographic Regions
• palearctic palearctic
  • native native
• ethiopian ethiopian
  • native native

Habitat

African golden wolves live in elevations from 0 to nearly 5,000 m. In the eastern part of their range, golden wolves primarily live in high elevations from 2,200 to 4,620 m. However, in the Sahara Desert, they can be found anywhere between sea level and 4,459 m, in isolated mountains, which are estimated to be refugia for golden wolves in the face of climate change.

Because of their generalist behavior and tolerance of dry habitats, golden wolves can be found in a wide range of habitats, including grasslands, coniferous temperate forests, invasive Eucalyptus forests, mangroves, dry plateaus, savannas, deserts, and semi-arid environments. However, their preferred habitat seems to be grasslands. Though golden wolves can occupy many habitats, the limiting factor seems to be access to water sources. In Tunisia, for example, differences in golden wolf distribution were best explained by the availability of water. In the Sahara Desert, golden wolves were found most frequently in areas with an annual rainfall of over 1,000 mm and were one of the few canids found in temperatures below 10°C. The only known unsuitable habitats seem to be extremely arid regions and dune fields.

Golden wolves do not occur solely in natural habitats: depending on the area, golden wolves can rely a great deal on human settlements, such as agricultural fields and rural areas. A study in northern Ethiopia found that golden wolf density can actually increase with increasing human density, rather than decreasing to avoid human activity, as is documented for many other canids. This is thought to occur because, in these areas, humans have depleted much of the native prey sources on which golden wolves rely. Therefore, golden wolves resort to feeding on human waste products in these areas. Golden wolves are thus urban exploiters, suggesting that golden wolves may thrive in the face of a growing human population.

• Habitat Regions
• temperate temperate
• tropical tropical
• terrestrial terrestrial

• Terrestrial Biomes
• desert or dune desert or dune
• savanna or grassland savanna or grassland
• forest forest
• mountains mountains

• Wetlands
• swamp swamp

• Other Habitat Features
• urban urban
• suburban suburban
• agricultural agricultural

Physical Description

African golden wolves were originally considered to be the same species as Eurasian golden jackals ( Canis aureus ). However, a study comparing mitochondrial DNA, microsatellites, sex chromosomes, and whole genomes showed that golden wolves have a unique gene pool from golden jackals and therefore constitute a different species. African wolves ( Canis lupaster ) were also concluded to be the same species as African golden wolves. This study also found that African golden wolves are more closely related to gray wolves ( Canis lupus ) than golden jackals. This close relation to gray wolves is interesting, considering that no gray wolves are found in Africa, but it is speculated that much of the canid diversity in Africa originated from Eurasian “wolf-like” colonizers that eventually went extinct. The first taxonomic description of an African golden wolf by Frédéric Cuvier in 1820 was also the first account that used the binomial nomenclature used for African golden wolves today: Canis anthus . Additionally, though Cuvier recognized African golden wolves as a separate species from golden jackals, the scientific community did not consider them as such until Koepfli et al.’s paper was published in 2015.

Golden wolves generally have an overall coat color of golden or pale yellow, dark tawny, or gray, depending on their local habitat. For example, the first golden wolf ever described lived in mountainous terrain and had a gray coat, whereas golden wolves in desert habitat tend to be golden. Regardless of overall coat color, most golden wolves have yellow markings and individual hairs with black, white, and tan bands. Their legs, tails, the back of their ears, and the top of their muzzles are all tan in color, and a back stripe runs down the upper third of their tails, which have black tips. Their throats, chests, stomachs, the inner sides of their legs, and the undersides of their jaws are white. Their hair is longest on their necks and tails, shortest on their heads and legs, and of intermediate length on the rest of their bodies. Hair runs from front to back across the whole body except between the front legs, where it instead runs back to front. Additional morphological characteristics include bushy tails, a thick under wool during winter, pale yellow to amber eyes, 7 to 8 mammary glands on females, and a dental formula of i 3/3, c 1/1, p 4/4, m 2/3.

Some sexual dimorphism occurs in golden wolves, mostly in terms of body size. Males have the same coloring and hair patterns as females, but are larger in size: males have a head and body length of 75 to 89.3 cm, a tail length of 20 to 34.7 cm, and weigh 6.3 to 15 kg. Females have a head and body length of 68 to 82.2 cm, a tail length of 20 to 29 cm, and weigh 6.5 to 10 kg. In general, females have 12% less body mass than males. There is also some sexual dimorphism in skull length in populations in East Africa, but not in North African populations.

Golden wolves in East Africa are smaller than golden wolves in West Africa, but otherwise look alike morphologically. There are also some small seasonal morphological differences, such as growing thick undercoats during winter and the emergence of a faint “black saddle” on their backs during some seasons. Overall, however, adult golden wolves look more or less the same, though individual identification may be possible by differences in the white markings on a their chests and throats. Even between age classes, golden wolves look very similar; adults can only be differentiated from juveniles by skeletal features. Adults have a high sagittal ridge on the front of their skulls that juveniles lack, and adults exhibit more tooth wear than young golden wolves.

Compared to Eurasian golden jackals ( Canis aureus ) - which scientists originally believed were the same species as golden wolves - African golden wolves look very similar in craniodental anatomy, size, and color. However, golden wolves have smaller muzzles and premolars, larger molars, and narrower, more pointed canines than golden jackals. Additionally, the entire lower third of the tails of golden jackals is black, whereas the tails of golden wolves are only black on their tips. The two species also differ geographically, as golden jackals are only found in Europe and golden wolves are only found in Africa.

On the eastern end of their range, golden wolves coexist with silver-backed jackals ( Canis mesomelas ) and side-striped jackals ( Canis adustus ), both of which are roughly the same size as African golden wolves. However, there are visible morphological differences between the three species: silver-backed jackals are easily identified by their red sides and legs and the silver “saddles” they have on their backs, and side-striped jackals have shorter ears, a gray stripe on their sides, and white-tipped tails. In comparison, golden wolves do not have a saddle, except for a vague black one in some seasons. They are also more gray and tan than red, do not have a side stripe, and have black-tipped tails.

• Sexual Dimorphism
• male larger

Reproduction

African golden wolves are monogamous, like many other canid species, though this can be somewhat flexible depending on the abundance of resources and shifting population characteristics. Pair bonds last for a lifetime, and a group will thus consist of the mated pair and their previous offspring, which help to raise young. When defending their territory, mated pairs will fight off intruders intrasexually: males attack other males and females attack other females. It is speculated that this may occur because an individual male wants to make sure he is the only one mating with his female, to ensure the pups he helps raise are his. Meanwhile, an individual female wants to make sure her male does not mate with other females, so that he will fully invest in helping to raise her pups. Thus, territoriality may help enforce monogamous pairs.

• Mating System
• monogamous monogamous

Both male and female golden wolves reach reproductive maturity at 10 to 11 months of age. Their breeding season is from October to December, with parturition occurring from December to March, sometimes even stretching on into April and May. Copulatory ties last for a few minutes, followed by a gestation period of about 63 days. Litter sizes can range from 1 to 9 pups, averaging around 6 pups per litter.

At birth, golden wolf pups are around 189 g. They are born blind and it takes 8 to 11 days for their eyes to open. Tooth eruption also occurs at 11 days. Golden wolf pups are born in underground dens, in which they stay until they are three weeks of age. Dens can have multiple openings, which are about 2 to 3 m long and about 1 m deep. Mothers stay in their dens with their pups and are supplemented by their mates. In some families, 11-to-18-year-old offspring from previous litters help raise new pups. The presence of these "helpers" has been shown to increase pup protection and provisioning, and thus survival. Both parents and their helpers assist in socializing the pups.

Even once pups emerge from their dens, they still rely on milk from their mother. However, they are also introduced to regurgitated food during this time. This food comes from their mothers and other adults in the group. Pups remain near their dens until they wean at 8 to 10 weeks of age. They begin assisting actively with foraging around 14 weeks of age. About 70% of pups will stay with their parents for up to two years and become helpers. During this time, they will not engage in breeding activity, even when they reach sexual maturity.

Only 30% of pups disperse before the next litter is born. It is speculated that most pups do not disperse right away because, due to high population densities at which golden wolves live, young pups would have a hard time finding a mate and establishing a territory of their own. However, once a juvenile does disperse from its family group, it is not yet known how it then finds a mate and establishes its own territory.

• Key Reproductive Features
• seasonal breeding seasonal breeding
• fertilization fertilization
• viviparous viviparous

As typical of canids, male golden wolf parental investment is high. Mated pairs are monogamous and raise their young together. Between birth and weaning, pups rely completely on the milk of their mothers. However, their fathers and previous offspring from the mating pair, will bring food back for mothers and defend dens. Both parents also assist in socializing the pups and, after the pups are weaned, regurgitate food for them.

• Parental Investment
• altricial altricial
• male parental care male parental care
• female parental care female parental care
• pre-hatching/birth
  • provisioning
    • male
    • female
  • protecting
    • male
    • female
• pre-weaning/fledging
  • provisioning
    • male
    • female
  • protecting
    • male
    • female
• pre-independence
  • provisioning
    • male
    • female
  • protecting
    • male
    • female
• post-independence association with parents

Lifespan/Longevity

The maximum known lifespan of an African golden wolf in the wild was observed to be 14 years; in captivity, the maximum lifespan is 18 years. However, usual lifespan in the wild ranges from 6 to 8 years, with an average lifespan of 7 years. Many golden wolves die as pups, as they are especially susceptible to disease and den flooding at this time. Little is known about what limits the lifespan of adults.

Behavior

African golden wolves are solitary, until they find a mate. Once a pair bond is formed, that pair stays together for life. The group size associated with a mating pair will grow and swell. This depends on how many pups they have and how many offspring stay to help raise the next batch of pups. In summary, golden wolves typically live in groups of two, though this number can increase with the presence of pups and helpers.

Strong intraspecific food competition usually selects against larger group sizes, though when food is abundant, large groups have been observed sharing a scavenged carcass.

Golden wolves are primarily diurnal - they are mostly active during the day, dawn, and dusk, and are not normally seen at night. Golden wolves are also highly mobile, with males seeming to move farther than females. One male was documented to move at least 230 km - with a high of 465 km - in 98 days. Additionally, in Tunisia there is high genetic diversity among golden wolves, suggesting that connectivity and dispersal capabilities are high between populations. Not much is known about how golden wolves find one another, including mates, or how they interact with non-family conspecifics.

• Key Behaviors
• cursorial
• diurnal diurnal
• crepuscular crepuscular
• nomadic nomadic
• sedentary sedentary
• solitary solitary
• territorial territorial
• social social

Home Range

Golden wolves are territorial and generally keep territories of about 0.39 to 5 km^2, though they have also been documented to stray past territorial borders in order to feed on carcasses. Territories are nestled within larger home ranges, the size of which depends on the age of individuals and the type of habitat.

Juveniles tend to have a much larger home range than adults, because of their need to spread out in search of mates and territory of their own. In woodland habitats, adult pairs have an average home range of 2.4 km^2, while dispersing juveniles have a much bigger home ranges, ranging from 5.6 to 21.7 km^2. Home range size in mountain habitats is much more variable, with adults having home ranges anywhere between 7.9 and 48.2 km^2 and dispersing juveniles having home ranges anywhere between 24.2 and 64.8 km^2. Both members of an adult pair will mark and defend their territory. They keep strict boundaries, though territories tend to overlap when individuals are part of a social group. Territories are generally held for about 8 years.

Communication and Perception

Scent markings and vocalizations are the primary ways that African golden wolves communicate with each other. These forms of communication are important for marking territory, mating, predator defense, and locating family members. These actions may be coupled with other signals, such as visual displays. For example, when an individual golden wolf marks its territory, it will urinate on specific landmarks with a raised leg, rather than in a squatting position, to show any golden wolves that may be watching that it is the holder of this particular territory.

African golden wolf vocalizations consist of howls, used for finding family members and asserting dominance, and growls and barks, which are used to warn family members of approaching predators. Barks are also used to stay in contact with group members during hunts for larger prey. Greeting ceremonies and grooming are also important ways of socializing.

• Communication Channels
• visual visual
• tactile tactile
• acoustic acoustic
• chemical chemical

• Other Communication Modes
• scent marks scent marks

• Perception Channels
• visual visual
• acoustic acoustic

Food Habits

African golden wolves primarily feed on wild boars ( Sus scrofa ) of all ages, though golden wolves likely only feed on adult boars as carrion, due to the dangers of actively hunting adult boars. Plant material also makes up a significant part of the diets of golden wolves. This includes various fruits, seeds, leaves and grasses for digestion and a source of water. Rabbits ( Oryctolagus cuniculus ) and livestock such as domestic sheep ( Ovis aries ) are also fairly common, as well as hares ( Lepus capensis ) and cats ( Felis lybicalcafus ) to a lesser extent. They have also been documented to eat birds (both wild and domesticated), rodents, and, more rarely, beetles. Thus, African golden wolves can be categorized as omnivores.

Adult pairs hunt together, but otherwise most golden wolves hunt alone. Individual golden wolves have been documented bringing down ungulates 4 to 5 times larger than themselves, though the success rate of mating pairs is higher than that of lone individuals. Adult pairs are also able to go after larger prey, such as Thomspon’s gazelle ( Eudorcus thomsonii ) and Abdim’s storks ( Ciconia abdimii ). If food is widely available, groups of up to 18 have been documented scavenging carcasses, but no documentation has been made of golden wolves hunting in large groups like their close relatives, gray wolves ( Canis lupus ).

To hunt rodents, golden wolves use their ears to pinpoint the exact location and either leap through the air to catch them or dig them out of their burrows. For ungulate prey, golden wolves generally focus on young, old, or injured individuals. They will chase these weaker individuals away from the rest of the herd, like many other canids. Golden wolves cache any leftovers for later. When a family group is on the hunt they will spread out rather than stay bunched together, with distances of a few hundred meters between each individual. They bark in order to stay in contact with one another during the hunt.

• Primary Diet
• omnivore omnivore

• Animal Foods
• birds
• mammals
• carrion carrion
• insects

• Plant Foods
• leaves
• seeds, grains, and nuts
• fruit

• Foraging Behavior
• stores or caches food stores or caches food

Predation

Spotted hyenas ( Crocuta crocuta ) are known to kill and eat African golden wolves in East Africa. Hyenas will often try to come into golden wolf dens to eat pups; when golden wolves see a hyena approaching their dens, they give a warning yowl, which alerts all of the adults nearby to chase the hyena away and bite its rump and genitals. Honey badgers ( Mellivora capensis ) have also been seen near golden wolf dens, but the adults have always chased them away before actual predation could be documented. Humans are also known to kill golden wolves in response to livestock predation. Besides aggressive actions by the adults guarding the den, not much is known about golden wolf anti-predator behavior.

Ecosystem Roles

Due to the widespread loss of many large carnivores in northern Africa, African golden wolves have become more or less the top predator. They are also shown to be opportunists and generalists, allowing them to spread widely across the landscape into many different ecosystems. One consequence of this is that they may be putting a large exploitative competition pressure on other predators, such as the common genet ( Genetta genetta ). Golden wolves are also speculated to compete exploitatively with fennec foxes ( Vulpes zerda ) and red foxes ( Vulpes Vulpes ) and appear to be the superior competitor, as foxes of both species have been shown to abandon water sources and hide whenever golden wolves approach. Because of this competition, it is believed that fennec foxes began to occupy more sandy areas that were less favorable to golden wolves and both fox species shifted to nocturnal activity to avoid golden wolves. Golden wolves are also known to have dietary overlap with black-backed jackals ( Canis mesomelas ) and side-striped jackals ( Canis adustus ) in East Africa. The degree to which this competition impacts these three species has yet to be documented.

Predation by golden wolves may help control rodent and boar populations. They are also scavengers, and thus are important for cycling energy and nutrients throughout their ecosystem. Highly mobile species such as golden wolves are especially important in providing these services across a wide range of systems. Thus, it is likely that golden wolves provide these vital ecosystem services.

Golden wolves seem to have a commensalistic relationship with cheetahs ( Acinonyx jubatus ), as documented in Kenya by Eaton (1969). When golden wolves encounter cheetahs, they will search the vicinity for a kill and, if they find one, scavenge off of it. If there is no kill immediately nearby, golden wolves will remain around the cheetahs for a while, following their movements until either a kill is made or the cheetahs remain inactive for too long and the golden wolves move on. Considering that golden wolves only feed on carcasses abandoned by cheetahs, this does not seem to be a parasitic relationship, as cheetahs are not prohibited from getting as much as they need to eat. There are also no records of cheetahs chasing golden wolves away from kills, suggesting that cheetahs are unaffected by their scavenging.

Additionally, cheetah and golden wolf family groups have been documented living near one another without fighting over resources or killing offspring. This suggests that there is little to no competitive relationship between the two species. In fact, it has been documented that, occasionally, golden wolves assist cheetah kills by distracting a herd while the cheetah sneaks up from behind, suggesting the relationship may be mutualistic. However, Eaton (1969) speculates that this behavior likely does not occur outside of their study area because of high competition between scavengers in other areas. Indeed, in Serengeti National Park, golden wolves are rarely observed on carcasses, and scavenged meat makes up only a small portion of their diet. This is thought to occur because of the competition with other scavengers and the danger posed to golden wolves by other scavengers and larger predators that made the kill.

Several golden wolves were shown to have antibodies for canine adenovirus, a liver infection, and canine coronavirus, a highly contagious intestinal disease. Both of these diseases can be easily spread to other canids through feces. Other individuals have tested positive for canine parvovirus, another intestinal disease that can spread to other canids, and canine distemper virus, a virus that affects the respiratory, gastrointestinal, and nervous systems. Canine distemper virus is an especially noteworthy disease because it can infect all sorts of other animals, including other canids, felids, and some primates. Other parasites include Coccidia , which are intestinal parasites that can affect canids and felids, as well as hookworms, tapeworms, mange, flukes, ticks, and Toxocara canis , another intestinal parasite that affects canids.

Economic Importance for Humans: Positive

It has been shown that organic waste from humans is a major food source for African golden wolves, which means they can assist with waste removal. In fact, they have been documented in northern Ethiopia, along with spotted hyenas ( Crocuta crocuta ), to remove organic waste that may be infected, therefore sanitizing rural areas. Studies of golden wolves have also given us a better understanding of how the domestication of dogs may have taken place.

• Positive Impacts
• research and education

Economic Importance for Humans: Negative

Similar to problems with gray wolves ( Canis lupus ) in the United States, predation by African golden wolves on livestock is a huge issue for rural communities in Africa. Golden wolves have been predating increasingly on livestock, and thus farmers have started retaliating. Between 2014 and 2015, farmers killed over 200 wolves. This is a serious problem in some areas, like Tunisia, where livestock has been documented to make up a significant part of the diets of golden wolves. However, relative frequency of livestock in their diet seems to primarily correlate with degree of livestock protection, suggesting that tighter management of livestock may be all that is required to solve this problem.

Additionally, golden wolves are hosts for the protozoan parasite Babesia gibsoni , which is commonly found in domestic dogs. Golden wolves are also reservoirs to Hepatozoon canis , as well as fleas. If golden wolves continue to become more frequent in human settlements, their presence could increase the spread of these parasites, and all of the other diseases mentioned above. These diseases could be spread to pets, other golden wolves and canids that congregate in human areas, and even livestock.

Some diseases that golden wolves carry, like flatworms, have been shown to also infect humans. Golden wolves are reservoirs to parasites like filarioids, which are responsible for pink eye and various lung diseases in humans, and guinea worms, which cause severe pain where the worm migrates as well as nausea and vomiting in humans. An increase of golden wolves in human settlements may lead to an increase in infections, which could be especially devastating for rural communities that may not have the means to treat them medically.

• Negative Impacts
• injures humans
  • carries human disease
• causes or carries domestic animal disease causes or carries domestic animal disease

Conservation Status

African golden wolves are listed as least concern but declining on the IUCN red list, and are not listed under the CITES appendices or the US Endangered Species Act. Durant et al. (2011) also documented a significant long-term decline in golden wolf populations in the Serengeti. Reasons for their decline may be due to overkill by hunters and poaching, both of which occur in the range of golden wolves, and retaliatory killing by farmers over predation of livestock. All of this is aided by the increased stock of automated weapons in places like Ethiopia. It is speculated that many golden wolves are also being affected by predator control programs for other species, primarily through the consumption of poisoned carcasses. Additionally, vehicular collisions were the source of death for at least fifty golden wolves in the Sahara Desert, which will likely have larger implications as countries develop and roads become more intricate and widespread.

Some countries in the range of golden wolves are currently or frequently in a state of war and other extreme conflict, which leads to increased habitat loss and fragmentation. Even the most remote regions, which usually experience little human presence, are affected by war, as opposing sides use these areas to gain tactical advantage. This pushes animals from places that may have once served as refugia. Because of this loss of habitat and refugia, many animals are locally extirpated. Additionally, the number of illegal killings were shown to increase drastically after a couple of years of war. The list of species killed likely includes golden wolves, because of their status as livestock predators. Outside of war-torn areas, widespread habitat loss due to human settlement, expansion, and over-grazing by livestock also occurs in the range of golden wolves. However, these are unlikely to have as large of an impact on golden wolf populations compared to the factors discussed above. Human settlements are less threatening to golden wolves likely because of how opportunistic they are, and due to their demonstrated ability to thrive in anthropogenic landscapes.

Part of the range of golden wolves is encompassed in the Tlemcen Hunting Reserve in Algeria. There are also several national parks scattered around Ethiopia and Eritrea, though many of these are poorly staffed and thus inadequately enforced. It is likely, however, that golden wolves receive at least some protection when considering all of the parks cumulatively.

Additional Links

Encyclopedia of Life

Contributors

Ken Feathers (author), University of Washington, Laura Prugh (editor), University of Washington, Galen Burrell (editor), Special Projects.

References

Amroun, M., P. Giraudoux, P. Delattre. 2006. A comparative study of the diets of two sympatric carnivores – the golden jackal (Canis aureus) and the common genet (Genetta genetta) – in Kabylia, Algeria. Mammalia , 70: 247-254.

Brito, J., A. Acosta, F. Álvares, F. Cuzin. 2009. Biogeography and conservation of taxa from remote regions: An application of ecological-niche based models and GIS to North-African canids. Biological Conservation , 142: 3020-3029.

Brito, J., S. Durant, N. Pettorelli, J. Newby, S. Canney, W. Algadafi, T. Rabeil, P. Crochet, J. Pleguezuelos, T. Wacher, K. de Smet, D. Gonçalves, M. da Silva, F. Martínez-Freiría, T. Abáigar, J. Campos, P. Comizzoli, S. Fahd, A. Fellous, H. Garba, D. Hamidou, A. Harouna, M. Hatcha, A. Nagy, T. Silva, A. Sow, C. Vale, Z. Boratyński, H. Rebelo, S. Carvalho. 2018. Armed conflicts and wildlife decline: Challenges and recommendations for effective conservation policy in the Sahara-Sahel. Conservation Letters , 11: 1-13.

Durant, S., M. Craft, R. Hilborn, S. Bashir, J. Hando, L. Thomas. 2011. Long-term trends in carnivore abundance using distance sampling in Serengeti National Park, Tanzania. Journal of Applied Ecology , 48: 1490-1500.

Eaton, R. 1969. Cooperative hunting by cheetahs and jackals and a theory of domestication of the dog. Mammalia , 33: 87-92.

Eddine, A., N. Mostefai, K. de Smet, D. Klees, H. Ansorge, Y. Karssene, C. Nowak, P. van der Leer. 2017. Diet composition of a newly recognized Canid species, the African golden wolf (Canis anthus), in Northern Algeria. Annales Zoologici Fennici , 54: 347-356.

Gherman, C., A. Mihalca. 2017. A synoptic overview of golden jackal parasites reveals high diversity of species. Parasites & Vectors , 10: 1-40.

Hunter, J., S. Durant, T. Caro. 2006. Patterns of scavenger arrival at cheetah kills in Serengeti National Park Tanzania. African Journal of Ecology , 45: 275-281.

Inger, R., E. Per, D. Cox, K. Gaston. 2016. Key role in ecosystem functioning of scavengers reliant on a single common species. Scientific Reports , 6: 1-5.

Karssene, Y., M. Chammem, F. Li, A. Eddine, A. Hermann, S. Nouira. 2019. Spatial and temporal variability in the distribution, daily activity and diet of fennec fox (Vulpes zerda), red fox (Vulpes vulpes) and African golden wolf (Canis anthus) in southern Tunisia. Mammalian Biology , 95: 41-50.

Karssene, Y., M. Chammem, C. Nowak, K. de Smet, D. Castro, A. Eddine, S. Lopes, V. Munoz-Fuentes, B. Cocchiararo, D. Klees, P. van der Leer, S. Nouira, R. Godinho. 2018. Noninvasive genetic assessment provides evidence of extensive gene flow and possible high movement ability in the African golden wolf. Mammalian Biology , 92: 94-101.

Koepfli, K., J. Pollinger, R. Godinho, J. Robinson, A. Lea, S. Hendricks, R. Schweizer, O. Thalmann, P. Silva, Z. Fan, A. Yurchenko, P. Dobrynin, A. Makunin, J. Cahill, B. Shapiro, F. Alvares, J. Brito, E. Geffen, J. Leonard, K. Helgen, W. Johnson, S. O'Brien, B. van Valkenburgh, R. Wayne. 2015. Genome-wide evidence reveals that African and Eurasian golden jackals are distinct species. Current Biology , 25: 2158-2165.

Maronpot, R., E. Guindy. 1970. Preliminary study of Babesia gibsoni Patton in wild carnivores and domesticated dogs in Egypt. American Journal of Veterinary Research , 31: 797-799.

Moehlman, P., V. Hayssen. 2018. Canis aureus (Carnivore: Canidae). Mammalian Species , 50: 14-25.

Moehlman, P. 1987. Social organization in jackals: The complex social system of jackals allows the successful rearing of very dependent young. American Scientist , 75: 366-375.

Tedford, R., X. Wang, B. Taylor. 2009. Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae). Bulletin of the American Museum of Natural History , 325: 1-218.

Yalden, D., M. Largen, D. Kock, J. Hillman. 1996. Catalogue of the mammals of Ethiopia and Eritrea. 7. Revised checklist, zoogeography and conservation. Tropical Zoology , 9: 73-164.

Yirga, G., H. Leirs, H. De Iongh, T. Asmelash, K. Gebrehiwot, M. Vos, H. Bauer. 2017. Densities of spotted hyaena (Crocuta crocuta) and African golden wolf (Canis anthus) increase with increasing anthropogenic influence. Mammalian Biology , 85: 60-69.