The Ethiopian wolf (Canis simensis), also called the red jackal, the Simien jackal or Simien fox, is a canine native to the Ethiopian Highlands, including the Simien Mountains. 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. 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. It is one of the world's rarest canids, and Africa's most endangered carnivore.

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.

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 the wolves through vaccination and community outreach programs.

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

Indigenous names

+Indigenous names for Canis simensis Linguistic group or area Indigenous name Literal translation Amharic ቀይ ቀበሮ (Ky kebero)ዋልጌ (Walgie) Red jackalTrickster Oromo Jeedala fardaaArouayé Horse's jackalReddish

Historical account

The species was first scientifically described in 1835 by Eduard Rüppell, who provided a skull for the British Museum. 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,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. while Charles Darwin hypothesised that the species gave rise to greyhounds.Darwin, Charles (1868). The Variation of Animals and Plants Under Domestication. Vol. I. Orange Judd. p. 48. 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 Powell-Cotton during his travels in Abyssinia.

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).

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.

Taxonomy and evolution

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. The authors of this study state that the ancestors of the Ethiopian wolf arrived in Africa from Eurasia at the same time as the ancestors of the African wild dog approximately 1.8 million years ago. The Ethiopian wolf has survived numerous climatic changes in its Ethiopian highland habitat, with its range repeatedly expanding and contracting with glacial cycles.

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.

See further: Canis evolution

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.

The Ethiopian wolf is one of two 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, and Oscar Neumann considered it to be "only an exaggerated fox". 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.

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. Further studies on RAD sequences found instances of Ethiopian wolves hybridizing with African golden wolves.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

Admixture with other Canis species 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 an extinct close 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.

Subspecies , two subspecies are recognised by Mammal Species of the World Volume Three (MSW3). SubspeciesTrinomial authorityDescriptionRangeSynonymsNorthern Ethiopian wolfC. s. simensis(Nominate subspecies)Rüppell, 1840Northwest Rift Valley: Simien Mountains, Mount Guna, Guassa Menz, north and south Wollo highlandsC. s. crinensis(Erlanger & Neumann, 1900)(Heuglin, 1862)C. s. simensis(Gray, 1869)C. s. walgi(Heuglin, 1862)Southern Ethiopian wolfC. s. citerniide Beaux, 1922This 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.Southeast Rift Valley: Arsi and Bale Mountains

Description

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 in body length, and in height. Adult males weigh , while females weigh .

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.

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

Behaviour

Social and territorial behaviours