Bat-eared fox
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The bat-eared fox (''Otocyon megalotis'') is a species of fox found on the African
savanna A savanna or savannah is a mixed woodland- grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground ...
. It is the only extant species of the genus ''Otocyon'' and considered a basal
canid Canidae (; from Latin, '' canis'', " dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found withi ...
species. Fossil records indicate this canid first appeared during the
middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. Th ...
. It is named for its large ears, which have a role in
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperatur ...
. The bat referred to in its colloquial name is possibly the Egyptian slit-faced bat (''Nycteris thebaica''), which is abundant in the region and has very large ears. Although not commonly used, other vernacular names include big-eared fox, black-eared fox, long-eared fox, Delalande's fox, cape fox, and motlosi.


Systematics and evolution

The bat-eared fox is the only living species of the genus ''Otocyon''. Its scientific name, given by
Anselme Gaëtan Desmarest Anselme Gaëtan Desmarest (6 March 1784 – 4 June 1838) was a French Zoology, zoologist and author. He was the son of Nicolas Desmarest and father of Eugène Anselme Sébastien Léon Desmarest. Desmarest was a disciple of Georges Cuvier and Alex ...
, was initially ''Canis megalotis'' (due to its close resemblance to jackals), and later changed by
Salomon Müller Salomon Müller (7 April 1804 – 29 December 1864) was a German naturalist. He was born in Heidelberg, and died in Freiburg im Breisgau. Müller was the son of a saddler in Heidelberg. Along with Heinrich Boie and Heinrich Christian Macklot, he w ...
which placed it in its own genus, ''Otocyon''; its huge ears and different dental formula warrant inclusion in a genus distinct from both ''Canis'' and true foxes (''Vulpes''). The generic name ''Otocyon'' is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words ''otus'' for ear and ''cyon'' for dog, while the specific name ''megalotis'' comes from the Greek words ''mega'' for large and ''otus'' for ear. Due to its different dentition, the bat-eared fox was previously placed in a distinct
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
of canids, Otocyoninae, as no relationship to any living species of canid could be established. However, according to more recent examinations, this species is regarded as having affinities with the vulpine line, and ''Otocyon'' was placed with high confidence as sister to the clade containing both the
raccoon dog The common raccoon dog (''Nyctereutes procyonoides''), also called the Chinese or Asian raccoon dog, is a small, heavy-set, fox-like canid native to East Asia. Named for its raccoon-like face markings, it is most closely related to foxes. Common ...
(''Nyctereutes'') and true foxes (''Vulpes''), occupying a basal position within
Canidae Canidae (; from Latin, '' canis'', " dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found withi ...
.


Subspecies

Currently, there are two recognized subspecies: * ''Otocyon megalotis megalotis'' (Desmarest, 1822) — found in southern Africa * ''Otocyon megalotis virgatus'' (Cabrera, 1910) — found in eastern Africa


Fossils

''Otocyon'' is poorly represented in the fossil record. It is suggested the genus forms a clade with ''
Prototocyon ''Prototocyon'' or ''Sivacyon'' is an extinct genus of small omnivorous canid living during the Late Pliocene and Early Pleistocene. Taxonomy ''Prototocyon'' was named by Pohle (1928) and was assigned to Canidae by Carroll (1988). Old literat ...
'', an extinct genus of canid. However, the generic distinction between ''Prototocyon'' and the extant ''Otocyon'' is doubtful. Fossils of an extinct fox-like canine known as ''Otocyon recki'', have been found in sediments of the
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropological localities in the world; the many sites exposed by the gorge have proven invaluable in furthering understanding of early human evolution. A steep-si ...
,
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
, dating back to the late Pliocene or early Pleistocene. However, it is now often placed in ''
Prototocyon ''Prototocyon'' or ''Sivacyon'' is an extinct genus of small omnivorous canid living during the Late Pliocene and Early Pleistocene. Taxonomy ''Prototocyon'' was named by Pohle (1928) and was assigned to Canidae by Carroll (1988). Old literat ...
''.


Description

Bat-eared foxes are relatively small canids, ranging in weight from 3 kg to 5.3 kg. Head and body length is 46–66 cm, tail length is 23–34 cm, shoulder height is 30–40 cm, and the notably large ears are 11–13 cm long. Generally, the pelage is tan-colored, with gray agouti guard hairs, giving its grizzled appearance, appearing more buff on the sides. The undersides and throat are pale. The limbs are dark, shading to dark brown or black at their extremities. The muzzle, the tip and upperside of the tail and the facial mask are black. The insides of the ears are white. Individuals of the East African subspecies, ''O. m. virgatus'', tend toward a buff pelage with dark brown markings, as opposed to the black of ''O. m. megalotis''. Proportionally large ears of bat-eared foxes, a characteristic shared by many other inhabitants of hot, arid climates, help to distribute heat. They also help in locating prey.


Range and distribution

The bat-eared fox has a disjointed range of distribution across the arid and semi-arid regions of Eastern and Southern Africa, in two
allopatric Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
populations (representing each of the recognized
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
) separated by approximately 1,000 km. Subspecies ''O. m. virgatus'' extends from southern
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,
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
and
Somalia Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constitut ...
, through
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The ...
and
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
to southwestern
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
; ''O. m. megalotis'' occurs in the southern part of Africa, ranging from
Angola , national_anthem = "Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordinat ...
through
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
and
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label= Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kal ...
to
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
, and extends as far east as
Mozambique Mozambique (), officially the Republic of Mozambique ( pt, Moçambique or , ; ny, Mozambiki; sw, Msumbiji; ts, Muzambhiki), is a country located in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi ...
and
Zimbabwe Zimbabwe (), officially the Republic of Zimbabwe, is a landlocked country located in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Botswana to the south-west, Zambia to the north, and ...
, spreading into the
Cape Peninsula The Cape Peninsula ( af, Kaapse Skiereiland) is a generally mountainous peninsula that juts out into the Atlantic Ocean at the south-western extremity of the African continent. At the southern end of the peninsula are Cape Point and the Ca ...
and toward
Cape Agulhas Cape Agulhas (; pt, Cabo das Agulhas , "Cape of the Needles") is a rocky headland in Western Cape, South Africa. It is the geographic southern tip of the African continent and the beginning of the dividing line between the Atlantic and Indian ...
. Home ranges vary in size from 0.3 to 3.5 km2. There are no confirmed observations in Zambia.


Habitat

Bat-eared foxes are adapted to arid or semi-arid environments. They are commonly found in short
grassland A grassland is an area where the vegetation is dominated by grasses ( Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur na ...
s, as well as the more arid regions of the savannas, along woodland edges, and in open acacia woodlands. They prefer bare ground and areas where grass is kept short by grazing
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
s and tend to hunt in these short grass and low shrub habitats. However, they do venture into areas with tall grasses and thick shrubs to hide when threatened. In addition to raising their young in dens, bat-eared foxes use self-dug dens for shelter from extreme temperatures and winds. They also lie under
acacia ''Acacia'', commonly known as the wattles or acacias, is a large genus of shrubs and trees in the subfamily Mimosoideae of the pea family Fabaceae. Initially, it comprised a group of plant species native to Africa and Australasia. The genus nam ...
trees in South Africa to seek shade during the day.


Diet

Bat-eared foxes are considered the only truly
insectivorous A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
canid Canidae (; from Latin, '' canis'', " dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found withi ...
, with a marked preference for harvester termites (''
Hodotermes mossambicus ''Hodotermes'' (from Greek ὁδός (hodós), travelling; Latin '' termes'', woodworm) is a genus of African harvester termites in the Hodotermitidae. They range from Palaearctic North Africa, through the East African savannas to the karroid ...
''), which can constitute 80–90% of its diet. When this particular species of termite is not available, their opportunistic diet allows a wide variety of food items to be taken: they can consume other species of termites, other
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s such as
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s,
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s (especially scarab beetles),
cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by st ...
s,
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
s,
millipede Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
s,
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s,
scorpion Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always en ...
s,
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s, and rarely birds, birds' eggs and chicks, small mammals, reptiles, and
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
(the desert truffle '' Kalaharituber pfeilii''). Berries, seeds, and wild fruit also are consumed. The bat-eared fox refuses to feed on snouted harvester termites, likely because it is not adapted to tolerate termites' chemical defense. Generally, bat-eared foxes meet their water requirements by the high water content of their diet. Water constitutes a critical resource during
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proces ...
.


Dentition

The teeth of the bat-eared fox are much smaller and reduced in shearing surface formation than teeth of other canid species. This is an adaptation to its
insectivorous A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
diet. The bat-eared fox is an old species that was widely distributed in the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
era. The teeth are not the bat-eared fox's only morphological adaptation for its diet. On the lower jaw, a step-like protrusion, called the subangular process, anchors the large
digastric muscle The digastric muscle (also digastricus) (named ''digastric'' as it has two 'bellies') is a small muscle located under the jaw. The term "digastric muscle" refers to this specific muscle. However, other muscles that have two separate muscle bell ...
to allow for rapid chewing. The
digastric muscle The digastric muscle (also digastricus) (named ''digastric'' as it has two 'bellies') is a small muscle located under the jaw. The term "digastric muscle" refers to this specific muscle. However, other muscles that have two separate muscle bell ...
is also modified to open and close the jaw five times per second.


Foraging

Bat-eared foxes usually hunt in groups, often splitting up in pairs, with separated subgroups moving through the same general area. When termites are plentiful, feeding aggregations of up to 15 individuals from different families occur. Individuals forage alone after family groups break in June or July and during the months after cub birth. Prey is located primarily by auditory means, rather than by smell or sight. Foraging patterns vary between seasons and populations, and coincide with termite availability. In eastern Africa, nocturnal foraging is the rule, while in southern Africa, nocturnal foraging during summer slowly changes to an almost solely diurnal pattern during the winter. Foraging techniques depend on prey type, but food is often located by walking slowly, nose close to the ground and ears tilted forward. It usually occurs in patches, which match the clumped prey resources, such as termite colonies, that also occur in patches. Groups are able to forage on clumps of prey in patches because they do not fight each other for food due to their degree of sociality and lack of territoriality.


Behavior

In the more northern areas of its range (around
Serengeti The Serengeti ( ) ecosystem is a geographical region in Africa, spanning northern Tanzania. The protected area within the region includes approximately of land, including the Serengeti National Park and several game reserves. The Serenget ...
), they are
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
85% of the time. However, around South Africa, they are nocturnal only in the summer and diurnal during the winter. Bat-eared foxes are highly social animals. They often live in pairs or groups, and home ranges of groups either overlap substantially or very little. In southern Africa, bat-eared foxes live in monogamous pairs with kits, while those in eastern Africa may live in pairs, or in stable family groups consisting of a male and up to three closely related females with cubs. Individuals forage, play, and rest together in a group, which helps in protection against predators. They engage in frequent and extended
allogrooming Social grooming is a behavior in which social animals, including humans, clean or maintain one another's body or appearance. A related term, allogrooming, indicates social grooming between members of the same species. Grooming is a major socia ...
sessions, which serve to strengthen group cohesion, mostly between mature adults, but also between young adults and mature adults Visual displays are very important in communication among bat-eared foxes. When they are looking intently at something, the head is held high, eyes are open, ears are erect and facing forward, and the mouth is closed. When an individual is in threat or showing submission, the ears are pulled back and lying against the head and the head is low. The tail also plays a role in communication. When an individual is asserting dominance or aggression, feeling threatened, playing, or being sexually aroused, the tail is arched in an inverted U shape. Individuals can also use
piloerection Goose bumps, goosebumps or goose-pimples are the bumps on a person's skin at the base of body hairs which may involuntarily develop when a person is tickled, cold or experiencing strong emotions such as fear, euphoria or sexual arousal. The f ...
, which occurs when individual hairs are standing straight, to make it appear larger when faced with extreme threat. When running, chasing, or fleeing, the tail is straight and horizontal. The bat-eared fox can recognize individuals up to 30 m away. The recognition process has three steps: First they ignore the individual, then they stare intently, and finally they either approach or attack without displays. When greeting another, the approaching individual shows symbolic submission which is received by the other individual with a high head and tail down. Few vocalizations are used for communication, but contact calls and warning calls are used, mostly during the winter. Glandular secretions and scratching, other than for digging, are absent in communication.


Reproduction

The bat-eared fox is predominantly socially
monogamous Monogamy ( ) is a form of dyadic relationship in which an individual has only one partner during their lifetime. Alternately, only one partner at any one time ( serial monogamy) — as compared to the various forms of non-monogamy (e.g., pol ...
, although it has been observed in
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
groups. In contrast to other canids, the bat-eared fox has a reversal in parental roles, with the male taking on the majority of the parental care behavior.
Gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pr ...
lasts for 60–70 days and females give birth to litters consisting of one to six kits. Beyond
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proces ...
, which lasts 14 to 15 weeks, males take over grooming, defending, huddling, chaperoning, and carrying the young between den sites. Additionally, male care and den attendance rates have been shown to have a direct correlation with cub survival rates. The female forages for food, which she uses to maintain milk production, on which the pups heavily depend. Food foraged by the female is not brought back to the pups or regurgitated to feed the pups. Pups in the
Kalahari The Kalahari Desert is a large semi-arid sandy savanna in Southern Africa extending for , covering much of Botswana, and parts of Namibia and South Africa. It is not to be confused with the Angolan, Namibian, and South African Namib coasta ...
region are born September–November and those in the
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label= Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kal ...
region are born October–December. Young bat-eared foxes disperse and leave their family groups at 5–6 months old and reach sexual maturity at 8–9 months. Bat-eared foxes have been recorded reaching maximum lifespans of over 14 to 17 years in captivity.


Conservation threats

The bat-eared fox has some commercial use for humans. They are important for
harvester termite The harvester termites (from Greek ὁδός (hodós), travelling; Latin '' termes'', woodworm) are an ancient, Old World family of termites, the Hodotermitidae. They are distinguished by the serrated inner edge of their mandibles, and their func ...
population control, as the termites are considered pests. They have also been hunted for their fur by Botswana natives. Additional threats to populations include disease and drought that can harm populations of prey; however, no major threats to bat-eared fox populations exist.


Notes


References

{{Taxonbar, from=Q200400
bat-eared fox The bat-eared fox (''Otocyon megalotis'') is a species of fox found on the African savanna. It is the only extant species of the genus ''Otocyon'' and considered a basal canid species. Fossil records indicate this canid first appeared during th ...
Carnivorans of Africa Fauna of East Africa Mammals of Southern Africa Mammals of Angola Mammals of Botswana Mammals of Ethiopia Mammals of Kenya Mammals of Namibia Mammals of South Africa Mammals of Tanzania Mammals of Zambia Myrmecophagous mammals
bat-eared fox The bat-eared fox (''Otocyon megalotis'') is a species of fox found on the African savanna. It is the only extant species of the genus ''Otocyon'' and considered a basal canid species. Fossil records indicate this canid first appeared during th ...