fbpx
Wikipedia

Ethiopian wolf

The Ethiopian wolf (Canis simensis),[3] also called the red jackal, the Simien jackal or Simien fox, is a canine native to the Ethiopian Highlands. In southeastern Ethiopia, it is also known as the horse jackal. It is similar to the coyote in size and build, and is distinguished by its long and narrow skull, and its red and white fur.[4] Unlike most large canids, which are widespread, generalist feeders, the Ethiopian wolf is a highly specialised feeder of Afroalpine rodents with very specific habitat requirements.[5] It is one of the world's rarest canids, and Africa's most endangered carnivore.[6]

Ethiopian wolf
Temporal range: Late Pleistocene – Recent
Ethiopian wolf on the Sanetti Plateau
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
C. simensis
Binomial name
Canis simensis
Ethiopian wolf range
Synonyms[3]
  • C. crinensis Erlanger and Neumann, 1900
  • C. semiensis Heuglin, 1862
  • C. simensis Gray, 1869
  • C. walgié Heuglin, 1862
  • C. citernii de Beaux, 1922

The species's current range is limited to seven isolated mountain ranges at altitudes of 3,000–4,500 m, with the overall adult population estimated at 360–440 individuals in 2011, more than half of them in the Bale Mountains.[1][7]

The Ethiopian wolf is listed as endangered by the IUCN, on account of its small numbers and fragmented range. Threats include increasing pressure from expanding human populations, resulting in habitat degradation through overgrazing, and disease transference and interbreeding from free-ranging dogs. Its conservation is headed by Oxford University's Ethiopian Wolf Conservation Programme, which seeks to protect wolves through vaccination and community outreach programs.[1]

Naming edit

Alternative English names for the Ethiopian wolf include the red jackal, the Simenian fox, the Simien jackal,[8] Ethiopian jackal, and Abyssinian wolf.[8]

Indigenous names edit

Historical account edit

 
Canis simensis by Friedrich Carl Vogel (from Eduard Rüppell's Neue Wirbelthiere zu der Fauna von Abyssinien gehörig).
 
Mounted specimen (1902), one of the first post-1835 specimens to reach Europe

The species was first scientifically described in 1835 by Eduard Rüppell,[12] who provided a skull for the British Museum.[13][14] European writers traveling in Ethiopia during the mid-19th century (then called Abyssinia) by Europeans and Ze Etiyopia by its citizens, wrote that the animal's skin was never worn by natives, as it was popularly believed that the wearer would die should any wolf hairs enter an open wound,[15] while Charles Darwin hypothesised that the species gave rise to greyhounds.[16][b] Since then, it was scarcely heard of in Europe up until the early 20th century, when several skins were shipped to England by Major Percy Horace Gordon Powell-Cotton during his travels in Abyssinia.[13][14]

The Ethiopian wolf was recognised as requiring protection in 1938, and received it in 1974. The first in-depth studies on the species occurred in the 1980s with the onset of the American-sponsored Bale Mountains Research Project. Ethiopian wolf populations in the Bale Mountains National Park were negatively affected by the political unrest of the Ethiopian Civil War, though the critical state of the species was revealed during the early 1990s after a combination of shooting and a severe rabies epidemic decimated most packs studied in the Web Valley and Sanetti Plateau. In response, the IUCN reclassified the species from endangered to critically endangered in 1994. The IUCN/SSC Canid Specialist Group advocated a three-front strategy of education, wolf population monitoring, and rabies control in domestic dogs. The establishment of the Ethiopian Wolf Conservation Programme in Bale soon followed in 1995 by Oxford University, in conjunction with the Ethiopian Wildlife Conservation Authority (EWCA).[6]

Soon after, a further wolf population was discovered in the Central Highlands. Elsewhere, information on Ethiopian wolves remained scarce; although first described in 1835 as living in the Simien Mountains, the paucity of information stemming from that area indicated that the species was likely declining there, while reports from the Gojjam plateau were a century out of date. Wolves were recorded in the Arsi Mountains since the early 20th century, and in the Bale Mountains in the late 1950s. The status of the Ethiopian wolf was reassessed in the late 1990s, following improvements in travel conditions into northern Ethiopia. The surveys taken revealed local extinctions in Mount Choqa, Gojjam, and in every northern Afroalpine region where agriculture is well developed and human pressure acute. This revelation stressed the importance of the Bale Mountains wolf populations for the species' long-term survival, as well as the need to protect other surviving populations. A decade after the rabies outbreak, the Bale populations had fully recovered to pre-epizootic levels, prompting the species' downlisting to endangered in 2004, though it still remains the world's rarest canid, and Africa's most endangered carnivore.[6]

Taxonomy and evolution edit

Phylogenetic tree of the wolf-like canids with timing in millions of years[c]
Caninae 3.5 Ma
3.0
2.5
2.0
0.96

Ethiopian wolf  

Dhole  

African wild dog  

The earliest known fossil of the Ethiopian wolf is known from the Melka Wakena paleoanthropological site-complex in the Southeastern Ethiopian Highlands. It is the right half of a mandible and dated to between 1.6 and 1.4 million years ago.[18]

In 1994, a mitochondrial DNA analysis showed a closer relationship to the gray wolf and the coyote than to other African canids, and C. simensis may be an evolutionary relic of a gray wolf-like ancestor's past invasion of northern Africa from Eurasia.[19]

See further: Canis evolution
 
Ethiopian wolf skull: Despite its close relation to the grey wolf, convergent evolution has resulted in a skull similar in shape to that of jackals and the South American maned wolf.[20]

Due to the high density of rodents in their new Afroalpine habitat, the ancestors of the Ethiopian wolf gradually developed into specialised rodent hunters. This specialisation is reflected in the animal's skull morphology, with its very elongated head, long jaw, and widely spaced teeth. During this period, the species likely attained its highest abundance, and had a relatively continuous distribution. This changed about 15,000 years ago with the onset of the current interglacial, which caused the species' Afroalpine habitat to fragment, thus isolating Ethiopian wolf populations from each other.[5]

The Ethiopian wolf is one of five Canis species present in Africa, and is readily distinguishable from jackals by its larger size, relatively longer legs, distinct reddish coat, and white markings. John Edward Gray and Glover Morrill Allen originally classified the species under a separate genus, Simenia,[21] and Oscar Neumann considered it to be "only an exaggerated fox".[22] Juliet Clutton-Brock refuted the separate genus in favour of placing the species in the genus Canis, upon noting cranial similarities with the side-striped jackal.[23]

In 2015, a study of mitochondrial genome sequences and whole genome nuclear sequences of African and Eurasian canids indicated that extant wolf-like canids have colonised Africa from Eurasia at least five times throughout the Pliocene and Pleistocene, which is consistent with fossil evidence suggesting that much of African canid fauna diversity resulted from the immigration of Eurasian ancestors, likely coincident with Plio-Pleistocene climatic oscillations between arid and humid conditions. According to a phylogeny derived from nuclear sequences, the Eurasian golden jackal (Canis aureus) diverged from the wolf/coyote lineage 1.9 million years ago, and with mitochondrial genome sequences indicating the Ethiopian wolf diverged from this lineage slightly prior to that.[24]: S1  Further studies on RAD sequences found instances of Ethiopian wolves hybridizing with African golden wolves.[25]

Admixture with other Canis species edit

In 2018, whole genome sequencing was used to compare members of the genus Canis. The study supports the African golden wolf being distinct from the golden jackal, and with the Ethiopian wolf being genetically basal to both. There are two genetically distinct African golden wolf populations that exist in northwestern and eastern Africa. This suggests that Ethiopian wolves – or a close and extinct relative – once had a much larger range within Africa to admix with other canids. There is evidence of gene flow between the eastern population and the Ethiopian wolf, which has led to the eastern population being distinct from the northwestern population. The common ancestor of both African golden wolf populations was a genetically admixed canid of 72% grey wolf and 28% Ethiopian wolf ancestry.[26]

Subspecies edit

As of 2005, two subspecies are recognised by Mammal Species of the World Volume Three (MSW3).[3]

Description edit

 
Painting (1926) by Louis Agassiz Fuertes

The Ethiopian wolf is similar in size and build to North America's coyote; it is larger than the black-backed jackal and side-striped jackals as well as the African wolf and has comparatively longer legs. Its skull is very flat, with a long facial region accounting for 58% of the skull's total length. The ears are broad, pointed, and directed forward. The teeth, particularly the premolars, are small and widely spaced. The canine teeth measure 14–22 mm in length, while the carnassials are relatively small. The Ethiopian wolf has eight mammae, of which only six are functional. The front paws have five toes, including a dewclaw, while the hind paws have four. As is typical in the genus Canis, males are larger than females, having 20% greater body mass. Adults measure 841–1,012 mm (33.1–39.8 in) in body length, and 530–620 mm (21–24 in) in height. Adult males weigh 14.2–19.3 kg (31–43 lb), while females weigh 11.2–14.15 kg (24.7–31.2 lb).[4]

The Ethiopian wolf has short guard hairs and thick underfur, which provides protection at temperatures as low as −15 °C. Its overall colour is ochre to rusty red, with dense whitish to pale ginger underfur. The fur of the throat, chest and underparts is white, with a distinct white band occurring around the sides of the neck. There is a sharp boundary between the red coat and white marks. The ears are thickly furred on the edges, though naked on the inside. The naked borders of the lips, the gums and palate are black. The lips, a small spot on the cheeks and an ascending crescent below the eyes are white. The thickly furred tail is white underneath, and has a black tip, though, unlike most other canids, there is no dark patch marking the supracaudal gland. It moults during the wet season (August–October), and there is no evident seasonal variation in coat colour, though the contrast between the red coat and white markings increases with age and social rank. Females tend to have paler coats than males. During the breeding season, the female's coat turns yellow, becomes woolier, and the tail turns brownish, losing much of its hair.[4]

Animals resulting from Ethiopian wolf-dog hybridisation tend to be more heavily built than pure wolves, and have shorter muzzles and different coat patterns.[27]

Behaviour edit

 
Southern Ethiopian wolf in the Bale Mountains

Social and territorial behaviours edit

The Ethiopian wolf is a social animal, living in family groups containing up to 20 adults (individuals older than one year), though packs of six wolves are more common. Packs are formed by dispersing males and a few females, which with the exception of the breeding female, are reproductively suppressed. Each pack has a well-established hierarchy, with dominance and subordination displays being common. Upon dying, a breeding female can be replaced by a resident daughter, though this increases the risk of inbreeding. Such a risk is sometimes circumvented by multiple paternity and extra-pack matings. The dispersal of wolves from their packs is largely restricted by the scarcity of unoccupied habitat.[28]

These packs live in communal territories, which encompass 6 km2 (2.3 sq mi) of land on average. In areas with little food, the species lives in pairs, sometimes accompanied by pups, and defends larger territories averaging 13.4 km2 (5.2 sq mi). In the absence of disease, Ethiopian wolf territories are largely stable, but packs can expand whenever the opportunity arises, such as when another pack disappears. The size of each territory correlates with the abundance of rodents, the number of wolves in a pack, and the survival of pups. Ethiopian wolves rest together in the open at night, and congregate for greetings and border patrols at dawn, noon, and evening. They may shelter from rain under overhanging rocks and behind boulders. The species never sleeps in dens, and only uses them for nursing pups. When patrolling their territories, Ethiopian wolves regularly scent-mark,[29] and interact aggressively and vocally with other packs. Such confrontations typically end with the retreat of the smaller group.[28]

Reproduction and development edit

The mating season usually takes place between August and November. Courtship involves the breeding male following the female closely. The breeding female only accepts the advances of the breeding male, or males from other packs. The gestation period is 60–62 days, with pups being born between October and December.[30] Pups are born toothless and with their eyes closed, and are covered in a charcoal-grey coat with a buff patch on the chest and abdomen. Litters consist of two to six pups, which emerge from their den after three weeks, when the dark coat is gradually replaced with the adult colouration. By the age of five weeks, the pups feed on a combination of milk and solid food, and become completely weaned off milk at the age of 10 weeks to six months.[4] All members of the pack contribute to protecting and feeding the pups, with subordinate females sometimes assisting the dominant female by suckling them. Full growth and sexual maturity are attained at the age of two years.[30] Cooperative breeding and pseudopregnancy have been observed in Ethiopian wolves.[31]

Most females disperse from their natal pack at about two years of age, and some become "floaters" that may successfully immigrate into existing packs. Breeding pairs are most often unrelated to each other, suggesting that female-biased dispersal reduces inbreeding.[32] Inbreeding is ordinarily avoided because it leads to a reduction in progeny fitness (inbreeding depression) due largely to the homozygous expression of deleterious recessive alleles.[33]

Hunting behaviours edit

 
Southern Ethiopian wolf feeding, Bale Mountains.

Unlike most social carnivores, the Ethiopian wolf tends to forage and feed on small prey alone. It is most active during the day, the time when rodents are themselves most active, though they have been observed to hunt in groups when targeting mountain nyala calves.[34] Major Percy-Cotton described the hunting behaviour of Ethiopian wolves as thus:

... they are most amusing to watch, when hunting. The rats, which are brown, with short tails, live in big colonies and dart from burrow to burrow, while the cuberow stands motionless till one of them shows, when he makes a pounce for it. If he is unsuccessful, he seems to lose his temper, and starts digging violently; but this is only lost labour, as the ground is honeycombed with holes, and every rat is yards away before he has thrown up a pawful.[35]

The technique described above is commonly used in hunting big-headed African mole-rats, with the level of effort varying from scratching lightly at the hole to totally destroying a set of burrows, leaving metre-high earth mounds.

Wolves in Bale have been observed to forage among cattle herds, a tactic thought to aid in ambushing rodents out of their holes by using the cattle to hide their presence.[4] Ethiopian wolves have also been observed forming temporary associations with troops of grazing geladas.[36] Solitary wolves hunt for rodents in the midst of the monkeys, ignoring juvenile monkeys, though these are similar in size to some of their prey. The monkeys, in turn, tolerate and largely ignore the wolves, although they take flight if they observe feral dogs, which sometimes prey on them. Within the troops, the wolves enjoy much higher success in capturing rodents than usual, perhaps because the monkeys' activities flush out the rodents, or because the presence of numerous larger animals makes it harder for rodents to spot a threat.[37]

Ecology edit

Habitat edit

 
Northern Ethiopian wolf in the Simien Mountains

The Ethiopian wolf is restricted to isolated pockets of Afroalpine grasslands and heathlands inhabited by Afroalpine rodents. Its ideal habitat extends from above the tree line around 3,200 to 4,500 m, with some wolves inhabiting the Bale Mountains being present in montane grasslands at 3,000 m. Although specimens were collected in Gojjam and northwestern Shoa at 2,500 m in the early 20th century, no recent records exist of the species occurring below 3,000 m. In modern times, subsistence agriculture, which extends up to 3,700 m, has largely restricted the species to the highest peaks.[38]

The Ethiopian wolf uses all Afroalpine habitats, but has a preference for open areas containing short herbaceous and grassland communities inhabited by rodents, which are most abundant along flat or gently sloping areas with poor drainage and deep soils. Prime wolf habitat in the Bale Mountains consists of short Alchemilla herbs and grasses, with low vegetation cover. Other favourable habitats consist of tussock grasslands, high-altitude scrubs rich in Helichrysum, and short grasslands growing in shallow soils. In its northern range, the wolf's habitat is composed of plant communities characterised by a matrix of Festuca tussocks, Euryops bushes, and giant lobelias, all of which are favoured by the wolf's rodent prey. Although marginal in importance, the ericaceous moorlands at 3,200–3,600 m in Simien may provide a refuge for wolves in highly disturbed areas.[38]

Diet edit

 
Big-headed mole rat (Tachyoryctes macrocephalus), one of the Ethiopian wolf's primary prey animals

In the Bale Mountains, the Ethiopian wolf's primary prey are big-headed African mole-rats, though it also feeds on grass rats, black-clawed brush-furred rats, and highland hares. Other secondary prey species include vlei rats, yellow-spotted brush-furred rats, and occasionally goslings and eggs. Ethiopian wolves have twice been observed to feed on rock hyraxes, and mountain nyala calves. It will also prey on reedbuck calves.[39] In areas where the big-headed African mole-rat is absent, the smaller Northeast African mole-rat is targeted.[39] In the Simien Mountains, the Ethiopian wolf preys on Abyssinian grass rats. Undigested sedge leaves have occasionally been found in Ethiopian wolf stomachs. The sedge possibly is ingested for roughage or for parasite control. The species may scavenge on carcasses, but is usually displaced by free-ranging dogs and African golden wolves. It typically poses no threat to livestock, with farmers often leaving herds in wolf-inhabited areas unattended.[4]

Range and populations edit

Six current Ethiopian wolf populations are known. North of the Rift Valley, the species occurs in the Simien Mountains in Gondar, in the northern and southern Wollo highlands, and in Guassa Menz in north Shoa. It has recently become extinct in Gosh Meda in north Shoa and Mount Guna, and has not been reported in Mount Choqa for several decades. Southeast of the Rift Valley, it occurs in the Arsi and Bale Mountains.[40]

Threats edit

The Ethiopian wolf has been considered rare since it was first recorded scientifically. The species likely has always been confined to Afroalpine habitats, so it was never widespread. In historical times, all of the Ethiopian wolf's threats are both directly and indirectly human-induced, as the wolf's highland habitat, with its high annual rainfall and rich fertile soils, is ideal for agricultural activities. Its proximate threats include habitat loss and fragmentation (subsistence agriculture, overgrazing, road construction, and livestock farming), diseases (primarily rabies and canine distemper), conflict with humans (poisoning, persecution, and road kills), and hybridisation with dogs.[42]

Disease edit

Rabies outbreaks, stemming from infected dogs, have killed many Ethiopian wolves over the 1990s and 2000s. Two well-documented outbreaks in Bale, one in 1991 and another in 2008–2009, resulted in the die-off or disappearance of 75% of known animals. Both incidents prompted reactive vaccinations in 2003 and 2008–2009, respectively. Canine distemper is not necessarily fatal to wolves, though a recent increase in infection has occurred, with outbreaks of canine distemper having been detected in 2005–2006 in Bale and in 2010 across subpopulations.[43]

Habitat loss edit

During the 1990s, wolf populations in Gosh Meda and Guguftu became extinct. In both cases, the extent of Afroalpine habitat above the limit of agriculture had been reduced to less than 20 km2. The EWCP team confirmed the extinction of a wolf population in Mt. Guna in 2011, whose numbers had been in single figures for several years. Habitat loss in the Ethiopian highlands is directly linked to agricultural expansion into Afroalpine areas. In the northern highlands, human density is among the highest in Africa, with 300 people per km2 in some localities, with almost all areas below 3,700 m having been converted into barley fields. Suitable areas of land below this limit are under some level of protection, such as Guassa-Menz and the Denkoro Reserve, or within the southern highlands, such as the Arsi and Bale Mountains. The most vulnerable wolf populations to habitat loss are those within relatively low-lying Afroalpine ranges, such as those in Aboi Gara and Delanta in North Wollo.[44]

Population fragmentation edit

Some Ethiopian wolf populations, particularly those in North Wollo, show signs of high fragmentation, which is likely to increase with current rates of human expansion. The dangers posed by fragmentation include increased contact with humans, dogs, and livestock, and further risk of isolation and inbreeding in wolf populations. Although no evidence of inbreeding depression or reduced fitness exists, the extremely small wolf population sizes, particularly those north of the Rift Valley, raise concerns among conservationists. Elsewhere, the Bale populations are fairly continuous, while those in Simien can still interbreed through habitat corridors.[45]

Encroachment within protected areas edit

In the Simien Mountains National Park, human and livestock populations are increasing by 2% annually, with further road construction allowing easy access to peasants into wolf home ranges; 3,171 people in 582 households were found to be living in the park and 1,477 outside the park in October 2005. Although the area of the park has since been expanded, further settlement stopped, and grazing restricted, effective enforcement may take years. As of 2011, about 30,000 people live in 30 villages around and two within the park, including 4,650 cereal farmers, herders, woodcutters, and many others. In Bale there are numerous villages in and around the area, comprising over 8,500 households with more than 12,500 dogs. In 2007, the estimate of households within wolf habitat numbered 1,756. Because of the high number of dogs, the risk of infection in local wolf populations is high. Furthermore, intentional and unintentional brush fires are frequent in the ericaceous moorlands wolves inhabit.[46]

Overgrazing edit

 
Ethiopian wolves resting alongside grazing zebu

Although wolves in Bale have learned to use cattle to conceal their presence when hunting for rodents, the level of grazing in the area can adversely affect the vegetation available for the wolves' prey. Although no declines in wolf populations related to overgrazing have occurred, high grazing intensities are known to lead to soil erosion and vegetation deterioration in Afroalpine areas such as Delanta and Simien.[47]

Human persecution and disturbance edit

Direct killings of wolves were more frequent during the Ethiopian Civil War, when firearms were more available. The extinction of wolves in Mt. Choqa was likely due to persecution. Although people living close to wolves in modern times believe that wolf populations are recovering, negative attitudes towards the species persist due to livestock predation. Wolves were largely unmolested by humans in Bale, as they were not considered threats to sheep and goats. However, they are perceived as threats to livestock elsewhere, with cases of retaliatory killings occurring in the Arsi Mountains. The Ethiopian wolf has not been recorded to be exploited for its fur, though in one case, wolf hides were used as saddle pads. It was once hunted by sportsmen, though this is now illegal. Vehicle collisions killed at least four wolves in the Sanetti Plateau since 1988, while two others were left with permanent limps. Similar accidents are a risk in areas where roads cut across wolf habitats, such as in Menz and Arsi.[27]

Hybridisation with dogs edit

Management plans for hybridization with dogs involve sterilization of known hybrids.[48] Incidences of Ethiopian wolf-dog hybridization have been recorded in Bale's Web Valley. At least four hybrids were identified and sterilized in the area. Although hybridization has not been detected elsewhere, scientists are concerned that it could pose a threat to the wolf population's genetic integrity, resulting in outbreeding depression or a reduction in fitness, though this does not appear to have taken place.[27] Due to the female's strong preference to avoid inbreeding, hybridization could be the result of not finding any males who are not close relatives outside of dogs.

Competition with African golden wolves edit

Encounters with African golden wolves (Canis lupaster) are usually agonistic, with Ethiopian wolves dominating African wolves if the latter enter their territories, and vice versa. Although African golden wolves are inefficient rodent hunters and thus not in direct competition with Ethiopian wolves, it is likely that heavy human persecution prevents the former from attaining numbers large enough to completely displace the latter.[49]

Conservation edit

The Ethiopian wolf is not listed on the CITES appendices, though it is afforded full official protection under Ethiopia's Wildlife Conservation Regulations of 1974, Schedule VI, with the killing of a wolf carrying a two-year jail sentence.[1]

The species is present in several protected areas, including three areas in South Wollo (Bale Mountains National Park, Simien Mountains National Park, and Borena Sayint Regional Park), one in north Shoa (Guassa Community Conservation Area), and one in the Arsi Mountains National Park. Areas of suitable wolf habitat have recently increased to 87%, as a result of boundary extensions in Simien and the creation of the Arsi Mountains National Park.[1]

Steps taken to ensure the survival of the Ethiopian wolf include dog vaccination campaigns in Bale, Menz, and Simien, sterilization programs for wolf-dog hybrids in Bale, rabies vaccination of wolves in parts of Bale, community and school education programs in Bale and Wollo, contributing to the running of national parks, and population monitoring and surveying. A 10-year national action plan was formed in February 2011.[1]

The species' critical situation was first publicised by the Wildlife Conservation Society in 1983, with the Bale Mountains Research Project being established shortly after. This was followed by a detailed, four-year field study, which prompted the IUCN/SSC Canid Specialist Group to produce an action plan in 1997. The plan called for the education of people in wolf-inhabited areas, wolf population monitoring, and the stemming of rabies in dog populations. The Ethiopian Wolf Conservation Programme was formed in 1995 by Oxford University, with donors including the Born Free Foundation, Frankfurt Zoological Society, and the Wildlife Conservation Network.[1]

The overall aim of the EWCP is to protect the wolf's Afroalpine habitat in Bale, and establish additional conservation areas in Menz and Wollo. The EWCP carries out education campaigns for people outside the wolf's range to improve dog husbandry and manage diseases within and around the park, as well as monitoring wolves in Bale, south and north Wollo. The program seeks to vaccinate up to 5,000 dogs a year to reduce rabies and distemper in wolf-inhabited areas.[1]

In 2016, the Korean company Sooam Biotech was reported to be attempting to clone the Ethiopian wolf using dogs as surrogate mothers to help conserve the species.[50]

Notes edit

  1. ^ This is in reference to the Ethiopian wolf's reported habit of following mares and cows about to give birth to feed on the afterbirth.[10]
  2. ^ This was later proven incorrect in 2010, when SNP studies showed that the dog's sole ancestor is the grey wolf.[17]
  3. ^ For a full set of supporting references refer to the note (a) in the phylotree at Evolution of the wolf#Wolf-like canids

References edit

  1. ^ a b c d e f g h Marino, J.; Sillero-Zubiri, C. (2011). "Canis simensis". IUCN Red List of Threatened Species. 2011: e.T3748A10051312. doi:10.2305/IUCN.UK.2011-1.RLTS.T3748A10051312.en. Retrieved 19 November 2021.
  2. ^ Rüppell 1835, Vol. Säugetiere: p. 39, pl. 14.
  3. ^ a b c Wozencraft, C. W. (2005). "Order Carnivora". In Wilson, D. E.; Reader, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference. Vol. 1 (3rd ed.). Johns Hopkins University Press. p. 577. ISBN 978-0-8018-8221-0.
  4. ^ a b c d e f g Sillero-Zubiri, C.; Gottelli, D. (1994). (PDF). Mammalian Species. 385 (485): 1–6. doi:10.2307/3504136. JSTOR 3504136. S2CID 253988540. Archived from the original (PDF) on 2015-09-24. Retrieved 2011-09-22.
  5. ^ a b Gottelli, D; Marino, J; Sillero-Zubiri, C; Funk, S (2004). (PDF). Molecular Ecology. 13 (8): 2275–2286. Bibcode:2004MolEc..13.2275G. doi:10.1111/j.1365-294x.2004.02226.x. PMID 15245401. S2CID 40027094. Archived from the original (PDF) on 2013-05-16.
  6. ^ a b c IUCN/SSC Canid Specialist Group 2011, pp. 7–8
  7. ^ "Ethiopian Wolf". Awf.org. Retrieved 19 March 2022.
  8. ^ a b Sillero-Zubiri, C.; Marino, J. (2004). "6.4 – Ethiopian wolf" (PDF). In Sillero-Zubiri, Claudio; Hoffmann, Michael; Macdonald, David Whyte (eds.). Canids: Foxes, Wolves, Jackals, and Dogs:Status Survey and Conservation Action Plan. IUCN-The World Conservation Union. pp. 167–174. ISBN 978-2831707860. Retrieved 4 July 2019.
  9. ^ Sillero-Zubiri & MacDonald 1997, p. 5 & 8
  10. ^ a b IUCN/SSC Canid Specialist Group 2011, p. 2
  11. ^ "Ethiopia | International Wolf Center". 28 July 2012.
  12. ^ Rüppell 1835, p. 39
  13. ^ a b Lydekker 1908, p. 462
  14. ^ a b Powell-Cotton 1902, pp. 206–207
  15. ^ Parkyns, Mansfield (1853). Life in Abyssinia: Being Notes Collected During Three Years' Residence and Travels in that Country. Vol. II. John Murray. pp. 12–13.
  16. ^ Darwin, Charles (1868). The Variation of Animals and Plants Under Domestication. Vol. I. Orange Judd. p. 48.
  17. ^ Vonholdt, B. M.; Pollinger, J. P.; Lohmueller, K. E.; Han, E.; Parker, H. G.; Quignon, P.; Degenhardt, J. D.; Boyko, A. R.; Earl, D. A.; Auton, A.; Reynolds, A.; Bryc, K.; Brisbin, A.; Knowles, J. C.; Mosher, D. S.; Spady, T. C.; Elkahloun, A.; Geffen, E.; Pilot, M.; Jedrzejewski, W.; Greco, C.; Randi, E.; Bannasch, D.; Wilton, A.; Shearman, J.; Musiani, M.; Cargill, M.; Jones, P. G.; Qian, Z.; Huang, W. (2010). "Genome-wide SNP and haplotype analyses reveal a rich history underlying dog domestication". Nature. 464 (7290): 898–902. Bibcode:2010Natur.464..898V. doi:10.1038/nature08837. PMC 3494089. PMID 20237475.
  18. ^ Martínez-Navarro, Bienvenido; Gossa, Tegenu; Carotenuto, Francesco; Bartolini-Lucenti, Saverio; Palmqvist, Paul; Asrat, Asfawossen; Figueirido, Borja; Rook, Lorenzo; Niespolo, Elizabeth M.; Renne, Paul R.; Herzlinger, Gadi; Hovers, Erella (2023-05-16). "The earliest Ethiopian wolf: implications for the species evolution and its future survival". Communications Biology. 6 (1): 530. doi:10.1038/s42003-023-04908-w. ISSN 2399-3642. PMC 10187515. PMID 37193884.
  19. ^ Gottelli, D.; Sillero-Zubiri, C.; Applebaum, G. D.; Roy, M. S.; Girman, D. J.; Garcia-Moreno, J.; Ostrander, E. A.; Wayne, R. K. (1994). "Molecular genetics of the most endangered canid: The Ethiopian wolf Canis simensis". Molecular Ecology. 3 (4): 301–12. Bibcode:1994MolEc...3..301G. doi:10.1111/j.1365-294X.1994.tb00070.x. PMID 7921357. S2CID 25976335.
  20. ^ Dalton, R. (2001). The skull morphology of the Ethiopian wolf (Canis simensis). B.Sc.thesis. University of Edinburgh, Edinburgh, UK.
  21. ^ IUCN/SSC Canid Specialist Group 2011, pp. 2–3
  22. ^ Powell-Cotton 1902, p. 459
  23. ^ Clutton-Brock, J.; Corbet, G.G.; Hills, M. (1976). "A review of the family Canidae, with a classification by numerical methods". Bull. Br. Mus. (Nat. Hist.). 29: 119–199. doi:10.5962/bhl.part.6922.
  24. ^ Koepfli, Klaus-Peter; Pollinger, John; Godinho, Raquel; Robinson, Jacqueline; Lea, Amanda; Hendricks, Sarah; Schweizer, Rena M.; Thalmann, Olaf; Silva, Pedro; Fan, Zhenxin; Yurchenko, Andrey A.; Dobrynin, Pavel; Makunin, Alexey; Cahill, James A.; Shapiro, Beth; Álvares, Francisco; Brito, José C.; Geffen, Eli; Leonard, Jennifer A.; Helgen, Kristofer M.; Johnson, Warren E.; o'Brien, Stephen J.; Van Valkenburgh, Blaire; Wayne, Robert K. (2015). "Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species". Current Biology. 25 (16): 2158–65. doi:10.1016/j.cub.2015.06.060. PMID 26234211.
  25. ^ Bahlk, S.H. (2015). Can hybridization be detected between African wolf and sympatric canids? . Master of Science Thesis. Center for Ecological and Evolutionary Synthesis Department of Bioscience Faculty of Mathematics and Natural Science, University of Oslo, Norway
  26. ^ Gopalakrishnan, Shyam; Sinding, Mikkel-Holger S.; Ramos-Madrigal, Jazmín; Niemann, Jonas; Samaniego Castruita, Jose A.; Vieira, Filipe G.; Carøe, Christian; Montero, Marc de Manuel; Kuderna, Lukas; Serres, Aitor; González-Basallote, Víctor Manuel; Liu, Yan-Hu; Wang, Guo-Dong; Marques-Bonet, Tomas; Mirarab, Siavash; Fernandes, Carlos; Gaubert, Philippe; Koepfli, Klaus-Peter; Budd, Jane; Rueness, Eli Knispel; Heide-Jørgensen, Mads Peter; Petersen, Bent; Sicheritz-Ponten, Thomas; Bachmann, Lutz; Wiig, Øystein; Hansen, Anders J.; Gilbert, M. Thomas P. (2018). "Interspecific Gene Flow Shaped the Evolution of the Genus Canis". Current Biology. 28 (21): 3441–3449.e5. doi:10.1016/j.cub.2018.08.041. PMC 6224481. PMID 30344120.
  27. ^ a b c IUCN/SSC Canid Specialist Group 2011, p. 32
  28. ^ a b IUCN/SSC Canid Specialist Group 2011, p. 4
  29. ^ Sillero-Zubiri, Claudio; Macdonald, David W. (1998). "Scent-marking and territorial behaviour of Ethiopian wolves Canis simensis" (PDF). Journal of Zoology. 245 (3): 351–361. doi:10.1017/s0952836998007134.
  30. ^ a b IUCN/SSC Canid Specialist Group 2011, pp. 4–5
  31. ^ van Kesteren, Freya; et al. (2013). "The physiology of cooperative breeding in a rare social canid; sex, suppression and pseudopregnancy in female Ethiopian wolves" (PDF). Physiology & Behavior. 122: 39–45. doi:10.1016/j.physbeh.2013.08.016. PMID 23994497. S2CID 46671897.
  32. ^ Randall, DA; Pollinger, JP; Wayne, RK; Tallents, LA; Johnson, PJ; Macdonald, DW (2007). "Inbreeding is reduced by female-biased dispersal and mating behavior in Ethiopian wolves". Behavioral Ecology. 18 (3): 579–89. doi:10.1093/beheco/arm010.
  33. ^ Charlesworth D, Willis JH (2009). "The genetics of inbreeding depression". Nat. Rev. Genet. 10 (11): 783–96. doi:10.1038/nrg2664. PMID 19834483. S2CID 771357.
  34. ^ IUCN/SSC Canid Specialist Group 2011, pp. 3–4
  35. ^ Powell-Cotton 1902, p. 207
  36. ^ Dartmouth College."Wolves are better hunters when monkeys are around: An unexpected co-existence in the Ethiopian highlands." ScienceDaily., 22 June 2015. <www.sciencedaily.com/releases/2015/06/150622085224.htm>.
  37. ^ Venkataraman, V. V.; Kerby, J. T.; Nguyen, N.; Ashenafi, Z. T.; Fashing, P. J. (2015-03-27). "Solitary Ethiopian wolves increase predation success on rodents when among grazing gelada monkey herds". Journal of Mammalogy. 96 (1): 129–137. doi:10.1093/jmammal/gyu013.
  38. ^ a b IUCN/SSC Canid Specialist Group 2011, pp. 19–20
  39. ^ a b "Canis simensis (Ethiopian wolf)". Animal Diversity Web.
  40. ^ IUCN/SSC Canid Specialist Group 2011, p. 10
  41. ^ IUCN/SSC Canid Specialist Group 2011, pp. 40–46
  42. ^ IUCN/SSC Canid Specialist Group 2011, p. 22
  43. ^ IUCN/SSC Canid Specialist Group 2011, pp. 22–26
  44. ^ IUCN/SSC Canid Specialist Group 2011, pp. 26–27
  45. ^ IUCN/SSC Canid Specialist Group 2011, pp. 27–28
  46. ^ IUCN/SSC Canid Specialist Group 2011, p. 29
  47. ^ IUCN/SSC Canid Specialist Group 2011, p. 30
  48. ^ Chris Barichievy; Shayne Clugston; Robert Sheldon. "Field report : Association between an Arabian wolf and a domestic dog in central Saudi Arabia" (PDF). Canids.org. Retrieved 19 March 2022.
  49. ^ Gutema, T. M., Foraging ecology and trophic niche overlap between sympatric African wolf and Ethiopian wolf in the Ethiopian Highlands 2016-10-06 at the Wayback Machine, The Rufford Foundation (November 2015)
  50. ^ Zastrow, Mark (8 February 2016). "Inside the cloning factory that creates 500 new animals a day". New Scientist. Retrieved 23 February 2016.

Cited works edit

  • IUCN; SSC Canid Specialist Group (2011). (PDF). IUCN/SSC Canid Specialist Group. Oxford, United Kingdom. Archived from the original (PDF) on 2014-09-08. Retrieved 2012-12-15.
  • Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited. {{cite journal}}: Cite journal requires |journal= (help)
  • Powell-Cotton, P. H. G. (1902). "A sporting trip through Abyssinia : a narrative of a nine months' journey from the plains of the Hawash to the snows of Simien, with a description of the game, from elephant to ibex, and notes on the manners and customs of the natives". London : Rowland Ward. {{cite journal}}: Cite journal requires |journal= (help)
  • Rüppell, Eduard (1835). Neue Wirbelthiere zu der Fauna von Abyssinien gehörig, entdeckt und beschrieben (in German). Frankfurt am Main : S. Schmerber.
  • Sillero-Zubiri, Claudio; MacDonald, David W. (1997). (PDF). IUCN/SSC Canid Specialist Group. ISBN 978-2-8317-0407-4. Archived from the original (PDF) on 26 December 2011. Retrieved 15 December 2012.

Further reading edit

  • Morell, V. 2006. . National Geographic Magazine, March 2006.
  • Williams, S. 2004. Ethiopian wolves on high. BBC Wildlife Magazine, July 2004.

External links edit

ethiopian, wolf, jackal, redirects, here, other, uses, jackal, disambiguation, canis, simensis, also, called, jackal, simien, jackal, simien, canine, native, ethiopian, highlands, southeastern, ethiopia, also, known, horse, jackal, similar, coyote, size, build. Red jackal redirects here For other uses see Red Jackal disambiguation The Ethiopian wolf Canis simensis 3 also called the red jackal the Simien jackal or Simien fox is a canine native to the Ethiopian Highlands In southeastern Ethiopia it is also known as the horse jackal It is similar to the coyote in size and build and is distinguished by its long and narrow skull and its red and white fur 4 Unlike most large canids which are widespread generalist feeders the Ethiopian wolf is a highly specialised feeder of Afroalpine rodents with very specific habitat requirements 5 It is one of the world s rarest canids and Africa s most endangered carnivore 6 Ethiopian wolfTemporal range Late Pleistocene RecentEthiopian wolf on the Sanetti PlateauConservation statusEndangered IUCN 3 1 1 Scientific classificationDomain EukaryotaKingdom AnimaliaPhylum ChordataClass MammaliaOrder CarnivoraFamily CanidaeGenus CanisSpecies C simensisBinomial nameCanis simensisRuppell 1840 2 Ethiopian wolf rangeSynonyms 3 C crinensis Erlanger and Neumann 1900 C semiensis Heuglin 1862 C simensis Gray 1869 C walgie Heuglin 1862 C citernii de Beaux 1922The species s current range is limited to seven isolated mountain ranges at altitudes of 3 000 4 500 m with the overall adult population estimated at 360 440 individuals in 2011 more than half of them in the Bale Mountains 1 7 The Ethiopian wolf is listed as endangered by the IUCN on account of its small numbers and fragmented range Threats include increasing pressure from expanding human populations resulting in habitat degradation through overgrazing and disease transference and interbreeding from free ranging dogs Its conservation is headed by Oxford University s Ethiopian Wolf Conservation Programme which seeks to protect wolves through vaccination and community outreach programs 1 Contents 1 Naming 1 1 Indigenous names 2 Historical account 3 Taxonomy and evolution 3 1 Admixture with other Canis species 3 2 Subspecies 4 Description 5 Behaviour 5 1 Social and territorial behaviours 5 2 Reproduction and development 5 3 Hunting behaviours 6 Ecology 6 1 Habitat 6 2 Diet 7 Range and populations 8 Threats 8 1 Disease 8 2 Habitat loss 8 3 Population fragmentation 8 4 Encroachment within protected areas 8 5 Overgrazing 8 6 Human persecution and disturbance 8 7 Hybridisation with dogs 8 8 Competition with African golden wolves 9 Conservation 10 Notes 11 References 12 Cited works 13 Further reading 14 External linksNaming editAlternative English names for the Ethiopian wolf include the red jackal the Simenian fox the Simien jackal 8 Ethiopian jackal and Abyssinian wolf 8 Indigenous names edit nbsp This section contains Ethiopic text Without proper rendering support you may see question marks boxes or other symbols instead of Ethiopic characters Indigenous names for Canis simensis 9 10 Linguistic group or area Indigenous name Literal translationAmharic ቀይ ቀበሮ Ky kebero ዋልጌ Walgie Red jackal 11 TricksterOromo Jeedala fardaaArouaye Horse s jackal a ReddishHistorical account edit nbsp Canis simensis by Friedrich Carl Vogel from Eduard Ruppell s Neue Wirbelthiere zu der Fauna von Abyssinien gehorig nbsp Mounted specimen 1902 one of the first post 1835 specimens to reach EuropeThe species was first scientifically described in 1835 by Eduard Ruppell 12 who provided a skull for the British Museum 13 14 European writers traveling in Ethiopia during the mid 19th century then called Abyssinia by Europeans and Ze Etiyopia by its citizens wrote that the animal s skin was never worn by natives as it was popularly believed that the wearer would die should any wolf hairs enter an open wound 15 while Charles Darwin hypothesised that the species gave rise to greyhounds 16 b Since then it was scarcely heard of in Europe up until the early 20th century when several skins were shipped to England by Major Percy Horace Gordon Powell Cotton during his travels in Abyssinia 13 14 The Ethiopian wolf was recognised as requiring protection in 1938 and received it in 1974 The first in depth studies on the species occurred in the 1980s with the onset of the American sponsored Bale Mountains Research Project Ethiopian wolf populations in the Bale Mountains National Park were negatively affected by the political unrest of the Ethiopian Civil War though the critical state of the species was revealed during the early 1990s after a combination of shooting and a severe rabies epidemic decimated most packs studied in the Web Valley and Sanetti Plateau In response the IUCN reclassified the species from endangered to critically endangered in 1994 The IUCN SSC Canid Specialist Group advocated a three front strategy of education wolf population monitoring and rabies control in domestic dogs The establishment of the Ethiopian Wolf Conservation Programme in Bale soon followed in 1995 by Oxford University in conjunction with the Ethiopian Wildlife Conservation Authority EWCA 6 Soon after a further wolf population was discovered in the Central Highlands Elsewhere information on Ethiopian wolves remained scarce although first described in 1835 as living in the Simien Mountains the paucity of information stemming from that area indicated that the species was likely declining there while reports from the Gojjam plateau were a century out of date Wolves were recorded in the Arsi Mountains since the early 20th century and in the Bale Mountains in the late 1950s The status of the Ethiopian wolf was reassessed in the late 1990s following improvements in travel conditions into northern Ethiopia The surveys taken revealed local extinctions in Mount Choqa Gojjam and in every northern Afroalpine region where agriculture is well developed and human pressure acute This revelation stressed the importance of the Bale Mountains wolf populations for the species long term survival as well as the need to protect other surviving populations A decade after the rabies outbreak the Bale populations had fully recovered to pre epizootic levels prompting the species downlisting to endangered in 2004 though it still remains the world s rarest canid and Africa s most endangered carnivore 6 Taxonomy and evolution editPhylogenetic tree of the wolf like canids with timing in millions of years c Caninae 3 5 Ma 3 0 2 5 2 0 0 96 0 6 0 38 Domestic dog nbsp Gray wolf nbsp Coyote nbsp African wolf nbsp Golden jackal nbsp Ethiopian wolf nbsp Dhole nbsp African wild dog nbsp 2 6 Side striped jackal nbsp Black backed jackal nbsp The earliest known fossil of the Ethiopian wolf is known from the Melka Wakena paleoanthropological site complex in the Southeastern Ethiopian Highlands It is the right half of a mandible and dated to between 1 6 and 1 4 million years ago 18 In 1994 a mitochondrial DNA analysis showed a closer relationship to the gray wolf and the coyote than to other African canids and C simensis may be an evolutionary relic of a gray wolf like ancestor s past invasion of northern Africa from Eurasia 19 See further Canis evolution nbsp Ethiopian wolf skull Despite its close relation to the grey wolf convergent evolution has resulted in a skull similar in shape to that of jackals and the South American maned wolf 20 Due to the high density of rodents in their new Afroalpine habitat the ancestors of the Ethiopian wolf gradually developed into specialised rodent hunters This specialisation is reflected in the animal s skull morphology with its very elongated head long jaw and widely spaced teeth During this period the species likely attained its highest abundance and had a relatively continuous distribution This changed about 15 000 years ago with the onset of the current interglacial which caused the species Afroalpine habitat to fragment thus isolating Ethiopian wolf populations from each other 5 The Ethiopian wolf is one of five Canis species present in Africa and is readily distinguishable from jackals by its larger size relatively longer legs distinct reddish coat and white markings John Edward Gray and Glover Morrill Allen originally classified the species under a separate genus Simenia 21 and Oscar Neumann considered it to be only an exaggerated fox 22 Juliet Clutton Brock refuted the separate genus in favour of placing the species in the genus Canis upon noting cranial similarities with the side striped jackal 23 In 2015 a study of mitochondrial genome sequences and whole genome nuclear sequences of African and Eurasian canids indicated that extant wolf like canids have colonised Africa from Eurasia at least five times throughout the Pliocene and Pleistocene which is consistent with fossil evidence suggesting that much of African canid fauna diversity resulted from the immigration of Eurasian ancestors likely coincident with Plio Pleistocene climatic oscillations between arid and humid conditions According to a phylogeny derived from nuclear sequences the Eurasian golden jackal Canis aureus diverged from the wolf coyote lineage 1 9 million years ago and with mitochondrial genome sequences indicating the Ethiopian wolf diverged from this lineage slightly prior to that 24 S1 Further studies on RAD sequences found instances of Ethiopian wolves hybridizing with African golden wolves 25 Admixture with other Canis species edit In 2018 whole genome sequencing was used to compare members of the genus Canis The study supports the African golden wolf being distinct from the golden jackal and with the Ethiopian wolf being genetically basal to both There are two genetically distinct African golden wolf populations that exist in northwestern and eastern Africa This suggests that Ethiopian wolves or a close and extinct relative once had a much larger range within Africa to admix with other canids There is evidence of gene flow between the eastern population and the Ethiopian wolf which has led to the eastern population being distinct from the northwestern population The common ancestor of both African golden wolf populations was a genetically admixed canid of 72 grey wolf and 28 Ethiopian wolf ancestry 26 Subspecies edit As of 2005 update two subspecies are recognised by Mammal Species of the World Volume Three MSW3 3 Subspecies Trinomial authority Description Range SynonymsNorthern Ethiopian wolfC s simensis Nominate subspecies nbsp Ruppell 1840 Northwest Rift Valley Simien Mountains Mount Guna Guassa Menz north and south Wollo highlands C s crinensis Erlanger amp Neumann 1900 C s semiensis Heuglin 1862 C s simensis Gray 1869 C s walgi Heuglin 1862 Southern Ethiopian wolfC s citernii nbsp de Beaux 1922 This canid was initially classed as a distinct subspecies on account of its bright red coat though this characteristic is unreliable as a taxonomic distinction However its nasal bones are consistently longer than those of the nominate subspecies 4 Southeast Rift Valley Arsi and Bale MountainsDescription edit nbsp Painting 1926 by Louis Agassiz FuertesThe Ethiopian wolf is similar in size and build to North America s coyote it is larger than the black backed jackal and side striped jackals as well as the African wolf and has comparatively longer legs Its skull is very flat with a long facial region accounting for 58 of the skull s total length The ears are broad pointed and directed forward The teeth particularly the premolars are small and widely spaced The canine teeth measure 14 22 mm in length while the carnassials are relatively small The Ethiopian wolf has eight mammae of which only six are functional The front paws have five toes including a dewclaw while the hind paws have four As is typical in the genus Canis males are larger than females having 20 greater body mass Adults measure 841 1 012 mm 33 1 39 8 in in body length and 530 620 mm 21 24 in in height Adult males weigh 14 2 19 3 kg 31 43 lb while females weigh 11 2 14 15 kg 24 7 31 2 lb 4 The Ethiopian wolf has short guard hairs and thick underfur which provides protection at temperatures as low as 15 C Its overall colour is ochre to rusty red with dense whitish to pale ginger underfur The fur of the throat chest and underparts is white with a distinct white band occurring around the sides of the neck There is a sharp boundary between the red coat and white marks The ears are thickly furred on the edges though naked on the inside The naked borders of the lips the gums and palate are black The lips a small spot on the cheeks and an ascending crescent below the eyes are white The thickly furred tail is white underneath and has a black tip though unlike most other canids there is no dark patch marking the supracaudal gland It moults during the wet season August October and there is no evident seasonal variation in coat colour though the contrast between the red coat and white markings increases with age and social rank Females tend to have paler coats than males During the breeding season the female s coat turns yellow becomes woolier and the tail turns brownish losing much of its hair 4 Animals resulting from Ethiopian wolf dog hybridisation tend to be more heavily built than pure wolves and have shorter muzzles and different coat patterns 27 Behaviour edit nbsp Southern Ethiopian wolf in the Bale MountainsSocial and territorial behaviours edit The Ethiopian wolf is a social animal living in family groups containing up to 20 adults individuals older than one year though packs of six wolves are more common Packs are formed by dispersing males and a few females which with the exception of the breeding female are reproductively suppressed Each pack has a well established hierarchy with dominance and subordination displays being common Upon dying a breeding female can be replaced by a resident daughter though this increases the risk of inbreeding Such a risk is sometimes circumvented by multiple paternity and extra pack matings The dispersal of wolves from their packs is largely restricted by the scarcity of unoccupied habitat 28 These packs live in communal territories which encompass 6 km2 2 3 sq mi of land on average In areas with little food the species lives in pairs sometimes accompanied by pups and defends larger territories averaging 13 4 km2 5 2 sq mi In the absence of disease Ethiopian wolf territories are largely stable but packs can expand whenever the opportunity arises such as when another pack disappears The size of each territory correlates with the abundance of rodents the number of wolves in a pack and the survival of pups Ethiopian wolves rest together in the open at night and congregate for greetings and border patrols at dawn noon and evening They may shelter from rain under overhanging rocks and behind boulders The species never sleeps in dens and only uses them for nursing pups When patrolling their territories Ethiopian wolves regularly scent mark 29 and interact aggressively and vocally with other packs Such confrontations typically end with the retreat of the smaller group 28 Reproduction and development edit The mating season usually takes place between August and November Courtship involves the breeding male following the female closely The breeding female only accepts the advances of the breeding male or males from other packs The gestation period is 60 62 days with pups being born between October and December 30 Pups are born toothless and with their eyes closed and are covered in a charcoal grey coat with a buff patch on the chest and abdomen Litters consist of two to six pups which emerge from their den after three weeks when the dark coat is gradually replaced with the adult colouration By the age of five weeks the pups feed on a combination of milk and solid food and become completely weaned off milk at the age of 10 weeks to six months 4 All members of the pack contribute to protecting and feeding the pups with subordinate females sometimes assisting the dominant female by suckling them Full growth and sexual maturity are attained at the age of two years 30 Cooperative breeding and pseudopregnancy have been observed in Ethiopian wolves 31 Most females disperse from their natal pack at about two years of age and some become floaters that may successfully immigrate into existing packs Breeding pairs are most often unrelated to each other suggesting that female biased dispersal reduces inbreeding 32 Inbreeding is ordinarily avoided because it leads to a reduction in progeny fitness inbreeding depression due largely to the homozygous expression of deleterious recessive alleles 33 Hunting behaviours edit nbsp Southern Ethiopian wolf feeding Bale Mountains Unlike most social carnivores the Ethiopian wolf tends to forage and feed on small prey alone It is most active during the day the time when rodents are themselves most active though they have been observed to hunt in groups when targeting mountain nyala calves 34 Major Percy Cotton described the hunting behaviour of Ethiopian wolves as thus they are most amusing to watch when hunting The rats which are brown with short tails live in big colonies and dart from burrow to burrow while the cuberow stands motionless till one of them shows when he makes a pounce for it If he is unsuccessful he seems to lose his temper and starts digging violently but this is only lost labour as the ground is honeycombed with holes and every rat is yards away before he has thrown up a pawful 35 The technique described above is commonly used in hunting big headed African mole rats with the level of effort varying from scratching lightly at the hole to totally destroying a set of burrows leaving metre high earth mounds Wolves in Bale have been observed to forage among cattle herds a tactic thought to aid in ambushing rodents out of their holes by using the cattle to hide their presence 4 Ethiopian wolves have also been observed forming temporary associations with troops of grazing geladas 36 Solitary wolves hunt for rodents in the midst of the monkeys ignoring juvenile monkeys though these are similar in size to some of their prey The monkeys in turn tolerate and largely ignore the wolves although they take flight if they observe feral dogs which sometimes prey on them Within the troops the wolves enjoy much higher success in capturing rodents than usual perhaps because the monkeys activities flush out the rodents or because the presence of numerous larger animals makes it harder for rodents to spot a threat 37 Ecology editHabitat edit nbsp Northern Ethiopian wolf in the Simien MountainsThe Ethiopian wolf is restricted to isolated pockets of Afroalpine grasslands and heathlands inhabited by Afroalpine rodents Its ideal habitat extends from above the tree line around 3 200 to 4 500 m with some wolves inhabiting the Bale Mountains being present in montane grasslands at 3 000 m Although specimens were collected in Gojjam and northwestern Shoa at 2 500 m in the early 20th century no recent records exist of the species occurring below 3 000 m In modern times subsistence agriculture which extends up to 3 700 m has largely restricted the species to the highest peaks 38 The Ethiopian wolf uses all Afroalpine habitats but has a preference for open areas containing short herbaceous and grassland communities inhabited by rodents which are most abundant along flat or gently sloping areas with poor drainage and deep soils Prime wolf habitat in the Bale Mountains consists of short Alchemilla herbs and grasses with low vegetation cover Other favourable habitats consist of tussock grasslands high altitude scrubs rich in Helichrysum and short grasslands growing in shallow soils In its northern range the wolf s habitat is composed of plant communities characterised by a matrix of Festuca tussocks Euryops bushes and giant lobelias all of which are favoured by the wolf s rodent prey Although marginal in importance the ericaceous moorlands at 3 200 3 600 m in Simien may provide a refuge for wolves in highly disturbed areas 38 Diet edit nbsp Big headed mole rat Tachyoryctes macrocephalus one of the Ethiopian wolf s primary prey animalsIn the Bale Mountains the Ethiopian wolf s primary prey are big headed African mole rats though it also feeds on grass rats black clawed brush furred rats and highland hares Other secondary prey species include vlei rats yellow spotted brush furred rats and occasionally goslings and eggs Ethiopian wolves have twice been observed to feed on rock hyraxes and mountain nyala calves It will also prey on reedbuck calves 39 In areas where the big headed African mole rat is absent the smaller Northeast African mole rat is targeted 39 In the Simien Mountains the Ethiopian wolf preys on Abyssinian grass rats Undigested sedge leaves have occasionally been found in Ethiopian wolf stomachs The sedge possibly is ingested for roughage or for parasite control The species may scavenge on carcasses but is usually displaced by free ranging dogs and African golden wolves It typically poses no threat to livestock with farmers often leaving herds in wolf inhabited areas unattended 4 Range and populations editSix current Ethiopian wolf populations are known North of the Rift Valley the species occurs in the Simien Mountains in Gondar in the northern and southern Wollo highlands and in Guassa Menz in north Shoa It has recently become extinct in Gosh Meda in north Shoa and Mount Guna and has not been reported in Mount Choqa for several decades Southeast of the Rift Valley it occurs in the Arsi and Bale Mountains 40 Summary of current status of Canis simensis 41 Area Habitat Population size estimates Status Importance Threats ConservationSimien Mountains North Gondar Patches connected by corridors totalling 273 km2 102 as of 2010 update Stable The second largest population the most genetically diverse is a tourist attraction Human disturbance road traffic extensive agriculture habitat degradation Helichrysum encroachment into rodent habitat disease and competition predation by golden wolves The entire range is within the Simien Mountains National Park and has been monitored regularly since 2003 Mt Guna South Gondar An isolated patch estimated at less than 20 km2 by 2004 As of 2011 update no sightings have been made despite intensive monitoring during two field visits Extinct Small population habitat loss isolation and possible competition with abundant golden wolf populations An ORDA Biodiversity Conservation Project is active in the area in conjunction with woreda and kebele governments and supported by the Guna Highland Water enterprise North Wollo Highlands Patchily distributed in an area of 140 km2 19 23 as of 2000 update Possibly declining otherwise stable as of 1998 Isolation habitat degradation human wildlife conflict and road encroachment The Frankfurt Zoological Society and its associates are working to create the Abuna Yoseph Community Conservation Area which is to encompass about a third of the wolf s range in North Wollo South Wollo highlands Patches connected by corridors totalling 243 km2 16 19 as of 2000 update Stable After Simien the second largest area north of the Rift Valley Overgrazing ploughing persecution and local negative attitudes The local ericaceous forests and grasslands are under the protection of the Borena Saiynt Regional Park from agriculture as low as 3 200 m The EWCP and the FZS have been involved in educational programs and wolf monitoring in the Denkoro area Guassa Menz North Shoa A single patch of 112 km2 As of 2010 update an estimated 40 have been missing since a canine distemper outbreak was detected in local dogs Although diminishing from disease the population is healthy and stable A core population with ideal habitat it is increasingly a tourist attraction Human disturbance rabies Helichrysum encroachment into rodent habitat and road traffic The wolf s range is protected by community resource management and the Guassa Community Conservation Area Educational campaigns are undertaken in schools near wolf ranges Arsi Mountains Bale 870 km2 54 wolves in 9 packs as of 2007 update 2010 Probably declining The third largest population in the second largest Afroalpine area in Ethiopia Habitat degradation expanding agriculture and road traffic Protected within the Arsi Mountains Regional ParkBale Mountains Bale 1 141 km2 About 250 adults and subadults Declining but stable in long term The largest population with the highest density of prey Disease rabies and distemper and agricultural expansion Most of the species habitat occurs within the Bale Mountains National Park Threats editThe Ethiopian wolf has been considered rare since it was first recorded scientifically The species likely has always been confined to Afroalpine habitats so it was never widespread In historical times all of the Ethiopian wolf s threats are both directly and indirectly human induced as the wolf s highland habitat with its high annual rainfall and rich fertile soils is ideal for agricultural activities Its proximate threats include habitat loss and fragmentation subsistence agriculture overgrazing road construction and livestock farming diseases primarily rabies and canine distemper conflict with humans poisoning persecution and road kills and hybridisation with dogs 42 Disease edit Rabies outbreaks stemming from infected dogs have killed many Ethiopian wolves over the 1990s and 2000s Two well documented outbreaks in Bale one in 1991 and another in 2008 2009 resulted in the die off or disappearance of 75 of known animals Both incidents prompted reactive vaccinations in 2003 and 2008 2009 respectively Canine distemper is not necessarily fatal to wolves though a recent increase in infection has occurred with outbreaks of canine distemper having been detected in 2005 2006 in Bale and in 2010 across subpopulations 43 Habitat loss edit During the 1990s wolf populations in Gosh Meda and Guguftu became extinct In both cases the extent of Afroalpine habitat above the limit of agriculture had been reduced to less than 20 km2 The EWCP team confirmed the extinction of a wolf population in Mt Guna in 2011 whose numbers had been in single figures for several years Habitat loss in the Ethiopian highlands is directly linked to agricultural expansion into Afroalpine areas In the northern highlands human density is among the highest in Africa with 300 people per km2 in some localities with almost all areas below 3 700 m having been converted into barley fields Suitable areas of land below this limit are under some level of protection such as Guassa Menz and the Denkoro Reserve or within the southern highlands such as the Arsi and Bale Mountains The most vulnerable wolf populations to habitat loss are those within relatively low lying Afroalpine ranges such as those in Aboi Gara and Delanta in North Wollo 44 Population fragmentation edit Some Ethiopian wolf populations particularly those in North Wollo show signs of high fragmentation which is likely to increase with current rates of human expansion The dangers posed by fragmentation include increased contact with humans dogs and livestock and further risk of isolation and inbreeding in wolf populations Although no evidence of inbreeding depression or reduced fitness exists the extremely small wolf population sizes particularly those north of the Rift Valley raise concerns among conservationists Elsewhere the Bale populations are fairly continuous while those in Simien can still interbreed through habitat corridors 45 Encroachment within protected areas edit In the Simien Mountains National Park human and livestock populations are increasing by 2 annually with further road construction allowing easy access to peasants into wolf home ranges 3 171 people in 582 households were found to be living in the park and 1 477 outside the park in October 2005 Although the area of the park has since been expanded further settlement stopped and grazing restricted effective enforcement may take years As of 2011 update about 30 000 people live in 30 villages around and two within the park including 4 650 cereal farmers herders woodcutters and many others In Bale there are numerous villages in and around the area comprising over 8 500 households with more than 12 500 dogs In 2007 the estimate of households within wolf habitat numbered 1 756 Because of the high number of dogs the risk of infection in local wolf populations is high Furthermore intentional and unintentional brush fires are frequent in the ericaceous moorlands wolves inhabit 46 Overgrazing edit nbsp Ethiopian wolves resting alongside grazing zebuAlthough wolves in Bale have learned to use cattle to conceal their presence when hunting for rodents the level of grazing in the area can adversely affect the vegetation available for the wolves prey Although no declines in wolf populations related to overgrazing have occurred high grazing intensities are known to lead to soil erosion and vegetation deterioration in Afroalpine areas such as Delanta and Simien 47 Human persecution and disturbance edit Direct killings of wolves were more frequent during the Ethiopian Civil War when firearms were more available The extinction of wolves in Mt Choqa was likely due to persecution Although people living close to wolves in modern times believe that wolf populations are recovering negative attitudes towards the species persist due to livestock predation Wolves were largely unmolested by humans in Bale as they were not considered threats to sheep and goats However they are perceived as threats to livestock elsewhere with cases of retaliatory killings occurring in the Arsi Mountains The Ethiopian wolf has not been recorded to be exploited for its fur though in one case wolf hides were used as saddle pads It was once hunted by sportsmen though this is now illegal Vehicle collisions killed at least four wolves in the Sanetti Plateau since 1988 while two others were left with permanent limps Similar accidents are a risk in areas where roads cut across wolf habitats such as in Menz and Arsi 27 Hybridisation with dogs edit Management plans for hybridization with dogs involve sterilization of known hybrids 48 Incidences of Ethiopian wolf dog hybridization have been recorded in Bale s Web Valley At least four hybrids were identified and sterilized in the area Although hybridization has not been detected elsewhere scientists are concerned that it could pose a threat to the wolf population s genetic integrity resulting in outbreeding depression or a reduction in fitness though this does not appear to have taken place 27 Due to the female s strong preference to avoid inbreeding hybridization could be the result of not finding any males who are not close relatives outside of dogs Competition with African golden wolves edit Encounters with African golden wolves Canis lupaster are usually agonistic with Ethiopian wolves dominating African wolves if the latter enter their territories and vice versa Although African golden wolves are inefficient rodent hunters and thus not in direct competition with Ethiopian wolves it is likely that heavy human persecution prevents the former from attaining numbers large enough to completely displace the latter 49 Conservation editThe Ethiopian wolf is not listed on the CITES appendices though it is afforded full official protection under Ethiopia s Wildlife Conservation Regulations of 1974 Schedule VI with the killing of a wolf carrying a two year jail sentence 1 The species is present in several protected areas including three areas in South Wollo Bale Mountains National Park Simien Mountains National Park and Borena Sayint Regional Park one in north Shoa Guassa Community Conservation Area and one in the Arsi Mountains National Park Areas of suitable wolf habitat have recently increased to 87 as a result of boundary extensions in Simien and the creation of the Arsi Mountains National Park 1 Steps taken to ensure the survival of the Ethiopian wolf include dog vaccination campaigns in Bale Menz and Simien sterilization programs for wolf dog hybrids in Bale rabies vaccination of wolves in parts of Bale community and school education programs in Bale and Wollo contributing to the running of national parks and population monitoring and surveying A 10 year national action plan was formed in February 2011 1 The species critical situation was first publicised by the Wildlife Conservation Society in 1983 with the Bale Mountains Research Project being established shortly after This was followed by a detailed four year field study which prompted the IUCN SSC Canid Specialist Group to produce an action plan in 1997 The plan called for the education of people in wolf inhabited areas wolf population monitoring and the stemming of rabies in dog populations The Ethiopian Wolf Conservation Programme was formed in 1995 by Oxford University with donors including the Born Free Foundation Frankfurt Zoological Society and the Wildlife Conservation Network 1 The overall aim of the EWCP is to protect the wolf s Afroalpine habitat in Bale and establish additional conservation areas in Menz and Wollo The EWCP carries out education campaigns for people outside the wolf s range to improve dog husbandry and manage diseases within and around the park as well as monitoring wolves in Bale south and north Wollo The program seeks to vaccinate up to 5 000 dogs a year to reduce rabies and distemper in wolf inhabited areas 1 In 2016 the Korean company Sooam Biotech was reported to be attempting to clone the Ethiopian wolf using dogs as surrogate mothers to help conserve the species 50 Notes edit This is in reference to the Ethiopian wolf s reported habit of following mares and cows about to give birth to feed on the afterbirth 10 This was later proven incorrect in 2010 when SNP studies showed that the dog s sole ancestor is the grey wolf 17 For a full set of supporting references refer to the note a in the phylotree at Evolution of the wolf Wolf like canidsReferences edit a b c d e f g h Marino J Sillero Zubiri C 2011 Canis simensis IUCN Red List of Threatened Species 2011 e T3748A10051312 doi 10 2305 IUCN UK 2011 1 RLTS T3748A10051312 en Retrieved 19 November 2021 Ruppell 1835 Vol Saugetiere p 39 pl 14 a b c Wozencraft C W 2005 Order Carnivora In Wilson D E Reader D M eds Mammal Species of the World A Taxonomic and Geographic Reference Vol 1 3rd ed Johns Hopkins University Press p 577 ISBN 978 0 8018 8221 0 a b c d e f g Sillero Zubiri C Gottelli D 1994 Canis simensis PDF Mammalian Species 385 485 1 6 doi 10 2307 3504136 JSTOR 3504136 S2CID 253988540 Archived from the original PDF on 2015 09 24 Retrieved 2011 09 22 a b Gottelli D Marino J Sillero Zubiri C Funk S 2004 The effect of the last glacial age on speciation and population genetic structure of the endangered Ethiopian wolf Canis simensis PDF Molecular Ecology 13 8 2275 2286 Bibcode 2004MolEc 13 2275G doi 10 1111 j 1365 294x 2004 02226 x PMID 15245401 S2CID 40027094 Archived from the original PDF on 2013 05 16 a b c IUCN SSC Canid Specialist Group 2011 pp 7 8 Ethiopian Wolf Awf org Retrieved 19 March 2022 a b Sillero Zubiri C Marino J 2004 6 4 Ethiopian wolf PDF In Sillero Zubiri Claudio Hoffmann Michael Macdonald David Whyte eds Canids Foxes Wolves Jackals and Dogs Status Survey and Conservation Action Plan IUCN The World Conservation Union pp 167 174 ISBN 978 2831707860 Retrieved 4 July 2019 Sillero Zubiri amp MacDonald 1997 p 5 amp 8 a b IUCN SSC Canid Specialist Group 2011 p 2 Ethiopia International Wolf Center 28 July 2012 Ruppell 1835 p 39 a b Lydekker 1908 p 462 a b Powell Cotton 1902 pp 206 207 Parkyns Mansfield 1853 Life in Abyssinia Being Notes Collected During Three Years Residence and Travels in that Country Vol II John Murray pp 12 13 Darwin Charles 1868 The Variation of Animals and Plants Under Domestication Vol I Orange Judd p 48 Vonholdt B M Pollinger J P Lohmueller K E Han E Parker H G Quignon P Degenhardt J D Boyko A R Earl D A Auton A Reynolds A Bryc K Brisbin A Knowles J C Mosher D S Spady T C Elkahloun A Geffen E Pilot M Jedrzejewski W Greco C Randi E Bannasch D Wilton A Shearman J Musiani M Cargill M Jones P G Qian Z Huang W 2010 Genome wide SNP and haplotype analyses reveal a rich history underlying dog domestication Nature 464 7290 898 902 Bibcode 2010Natur 464 898V doi 10 1038 nature08837 PMC 3494089 PMID 20237475 Martinez Navarro Bienvenido Gossa Tegenu Carotenuto Francesco Bartolini Lucenti Saverio Palmqvist Paul Asrat Asfawossen Figueirido Borja Rook Lorenzo Niespolo Elizabeth M Renne Paul R Herzlinger Gadi Hovers Erella 2023 05 16 The earliest Ethiopian wolf implications for the species evolution and its future survival Communications Biology 6 1 530 doi 10 1038 s42003 023 04908 w ISSN 2399 3642 PMC 10187515 PMID 37193884 Gottelli D Sillero Zubiri C Applebaum G D Roy M S Girman D J Garcia Moreno J Ostrander E A Wayne R K 1994 Molecular genetics of the most endangered canid The Ethiopian wolf Canis simensis Molecular Ecology 3 4 301 12 Bibcode 1994MolEc 3 301G doi 10 1111 j 1365 294X 1994 tb00070 x PMID 7921357 S2CID 25976335 Dalton R 2001 The skull morphology of the Ethiopian wolf Canis simensis B Sc thesis University of Edinburgh Edinburgh UK IUCN SSC Canid Specialist Group 2011 pp 2 3 Powell Cotton 1902 p 459 Clutton Brock J Corbet G G Hills M 1976 A review of the family Canidae with a classification by numerical methods Bull Br Mus Nat Hist 29 119 199 doi 10 5962 bhl part 6922 Koepfli Klaus Peter Pollinger John Godinho Raquel Robinson Jacqueline Lea Amanda Hendricks Sarah Schweizer Rena M Thalmann Olaf Silva Pedro Fan Zhenxin Yurchenko Andrey A Dobrynin Pavel Makunin Alexey Cahill James A Shapiro Beth Alvares Francisco Brito Jose C Geffen Eli Leonard Jennifer A Helgen Kristofer M Johnson Warren E o Brien Stephen J Van Valkenburgh Blaire Wayne Robert K 2015 Genome wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species Current Biology 25 16 2158 65 doi 10 1016 j cub 2015 06 060 PMID 26234211 Bahlk S H 2015 Can hybridization be detected between African wolf and sympatric canids Master of Science Thesis Center for Ecological and Evolutionary Synthesis Department of Bioscience Faculty of Mathematics and Natural Science University of Oslo Norway Gopalakrishnan Shyam Sinding Mikkel Holger S Ramos Madrigal Jazmin Niemann Jonas Samaniego Castruita Jose A Vieira Filipe G Caroe Christian Montero Marc de Manuel Kuderna Lukas Serres Aitor Gonzalez Basallote Victor Manuel Liu Yan Hu Wang Guo Dong Marques Bonet Tomas Mirarab Siavash Fernandes Carlos Gaubert Philippe Koepfli Klaus Peter Budd Jane Rueness Eli Knispel Heide Jorgensen Mads Peter Petersen Bent Sicheritz Ponten Thomas Bachmann Lutz Wiig Oystein Hansen Anders J Gilbert M Thomas P 2018 Interspecific Gene Flow Shaped the Evolution of the Genus Canis Current Biology 28 21 3441 3449 e5 doi 10 1016 j cub 2018 08 041 PMC 6224481 PMID 30344120 a b c IUCN SSC Canid Specialist Group 2011 p 32 a b IUCN SSC Canid Specialist Group 2011 p 4 Sillero Zubiri Claudio Macdonald David W 1998 Scent marking and territorial behaviour of Ethiopian wolves Canis simensis PDF Journal of Zoology 245 3 351 361 doi 10 1017 s0952836998007134 a b IUCN SSC Canid Specialist Group 2011 pp 4 5 van Kesteren Freya et al 2013 The physiology of cooperative breeding in a rare social canid sex suppression and pseudopregnancy in female Ethiopian wolves PDF Physiology amp Behavior 122 39 45 doi 10 1016 j physbeh 2013 08 016 PMID 23994497 S2CID 46671897 Randall DA Pollinger JP Wayne RK Tallents LA Johnson PJ Macdonald DW 2007 Inbreeding is reduced by female biased dispersal and mating behavior in Ethiopian wolves Behavioral Ecology 18 3 579 89 doi 10 1093 beheco arm010 Charlesworth D Willis JH 2009 The genetics of inbreeding depression Nat Rev Genet 10 11 783 96 doi 10 1038 nrg2664 PMID 19834483 S2CID 771357 IUCN SSC Canid Specialist Group 2011 pp 3 4 Powell Cotton 1902 p 207 Dartmouth College Wolves are better hunters when monkeys are around An unexpected co existence in the Ethiopian highlands ScienceDaily 22 June 2015 lt www sciencedaily com releases 2015 06 150622085224 htm gt Venkataraman V V Kerby J T Nguyen N Ashenafi Z T Fashing P J 2015 03 27 Solitary Ethiopian wolves increase predation success on rodents when among grazing gelada monkey herds Journal of Mammalogy 96 1 129 137 doi 10 1093 jmammal gyu013 a b IUCN SSC Canid Specialist Group 2011 pp 19 20 a b Canis simensis Ethiopian wolf Animal Diversity Web IUCN SSC Canid Specialist Group 2011 p 10 IUCN SSC Canid Specialist Group 2011 pp 40 46 IUCN SSC Canid Specialist Group 2011 p 22 IUCN SSC Canid Specialist Group 2011 pp 22 26 IUCN SSC Canid Specialist Group 2011 pp 26 27 IUCN SSC Canid Specialist Group 2011 pp 27 28 IUCN SSC Canid Specialist Group 2011 p 29 IUCN SSC Canid Specialist Group 2011 p 30 Chris Barichievy Shayne Clugston Robert Sheldon Field report Association between an Arabian wolf and a domestic dog in central Saudi Arabia PDF Canids org Retrieved 19 March 2022 Gutema T M Foraging ecology and trophic niche overlap between sympatric African wolf and Ethiopian wolf in the Ethiopian Highlands Archived 2016 10 06 at the Wayback Machine The Rufford Foundation November 2015 Zastrow Mark 8 February 2016 Inside the cloning factory that creates 500 new animals a day New Scientist Retrieved 23 February 2016 Cited works editIUCN SSC Canid Specialist Group 2011 Strategic Planning for Ethiopian Wolf Conservation PDF IUCN SSC Canid Specialist Group Oxford United Kingdom Archived from the original PDF on 2014 09 08 Retrieved 2012 12 15 Lydekker Richard 1908 The Game Animals of Africa London R Ward limited a href Template Cite journal html title Template Cite journal cite journal a Cite journal requires journal help Powell Cotton P H G 1902 A sporting trip through Abyssinia a narrative of a nine months journey from the plains of the Hawash to the snows of Simien with a description of the game from elephant to ibex and notes on the manners and customs of the natives London Rowland Ward a href Template Cite journal html title Template Cite journal cite journal a Cite journal requires journal help Ruppell Eduard 1835 Neue Wirbelthiere zu der Fauna von Abyssinien gehorig entdeckt und beschrieben in German Frankfurt am Main S Schmerber Sillero Zubiri Claudio MacDonald David W 1997 The Ethiopian Wolf Status Survey and Conservation Action Plan PDF IUCN SSC Canid Specialist Group ISBN 978 2 8317 0407 4 Archived from the original PDF on 26 December 2011 Retrieved 15 December 2012 Further reading editMorell V 2006 Wolves of Ethiopia National Geographic Magazine March 2006 Williams S 2004 Ethiopian wolves on high BBC Wildlife Magazine July 2004 External links edit nbsp Wikimedia Commons has media related to Canis simensis nbsp Wikispecies has information related to Canis simensis Ethiopian Wolf Conservation Programme EWCP Conservation of Ethiopian wolves Canis simensis at WildCRU of the University of Oxford Department of Zoology Ethiopian Wolf Canis simensis from the IUCN Species Survival Commission Canid Specialist Group of the International Union for Conservation of Nature Supporting the EWCP Archived 2009 02 12 at the Wayback Machine at the Born Free Foundation Ethiopian wolf Canis simensis at ARKive images and videos Retrieved from https en wikipedia org w index php title Ethiopian wolf amp oldid 1207378821, wikipedia, wiki, book, books, library,

article

, read, download, free, free download, mp3, video, mp4, 3gp, jpg, jpeg, gif, png, picture, music, song, movie, book, game, games.