Aardwolf

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The aardwolf (Proteles cristatus<ref>Template:Cite journal</ref>) is an insectivorous hyaenid species, native to East and Southern Africa. Its name means "earth-wolf" in Afrikaans and Dutch.<ref name=EB/><ref>"Aardwolf, n." Dictionary of South African English. Dictionary Unit for South African English, 2018. Web. 25 February 2019.</ref> It is also called the maanhaar-jackal<ref name="OED">Template:Harvnb</ref><ref>"Maanhaar, n." Dictionary of South African English. Dictionary Unit for South African English, 2018. Web. 25 February 2019.</ref> (Afrikaans for "mane-jackal"), termite-eating hyena<ref>Template:Cite book</ref> and civet hyena, based on its habit of secreting substances from its anal gland, a characteristic shared with the African civet.<ref name=ingo/>

Unlike many of its relatives in the order Carnivora, the aardwolf does not hunt large animals. It eats insects and their larvae,<ref name="wff">Template:Harvnb</ref> mainly termites; one aardwolf can lap up as many as 300,000 termites during a single nightTemplate:Sfn using its long, sticky tongue. The aardwolf's tongue has adapted to be tough enough to withstand the strong bite of termites.<ref name=mh71/>

The aardwolf lives in the shrublands of eastern and southern Africa – open lands covered with stunted trees and shrubs. It is nocturnal, resting in burrows during the day and emerging at night to seek food.

TaxonomyEdit

The aardwolf is generally classified as part of the hyena family Hyaenidae. However, it was formerly placed in its own family Protelidae.Template:Refn Early on, scientists felt that it was merely mimicking the striped hyena, which subsequently led to the creation of Protelidae.<ref>Template:Harvnb</ref> Recent studies have suggested that the aardwolf probably diverged from other hyaenids early on; how early is still unclear, as the fossil record and genetic studies disagree by 10 million years.<ref name=koe>Template:Harvnb</ref>Template:Refn

The aardwolf is the only surviving species in the subfamily Protelinae. There is disagreement as to whether the species is monotypic,<ref name=Woz>Template:Harvnb</ref> or can be divided into subspecies. A 2021 study found the genetic differences in eastern and southern aardwolves may be pronounced enough to categorize them as species.<ref>Template:Cite journal</ref>

Image	Subspecies	Distribution
File:Aardwolf, Proteles cristata, at Lion and Rhino Reserve, Gauteng, South Africa (47987215058).jpg	P. c. cristatus	Southern Africa
File:Aardwolf, Buffalo Springs National Park, Kenya 1 (cropped).jpg	P. c. septentrionalis Template:Au	East Africa.<ref name=ingo/><ref name=mh33/>

A 2006 molecular analysis indicates it is phylogenetically the most basal of the four extant hyaenidae species.Template:Sfn

Template:Clade

EtymologyEdit

The generic name Proteles is derived from two words of Greek origin: Template:Langx, prōtos and Template:Langx téleios, which combined mean "complete in front" referring to the aardwolf's five toes on the front paws and four on the hindpaws.<ref name=ingo/> The specific name cristatus is derived from Latin and means "provided with a comb or tuft", relating to its mane.<ref name=ingo/>

DescriptionEdit

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File:Aardwolf, Proteles cristata, at Lion and Rhino Reserve, Gauteng, South Africa (47987248651).jpg

Aardwolf with tongue exposed. Note the prominent papillae.

The aardwolf resembles a much smaller and thinner striped hyena, with a more slender muzzle, black vertical stripes on a coat of yellowish fur, and a long, distinct mane down the midline of the neck and back. It also has one or two diagonal stripes down the fore and hindquarters and several stripes on its legs.<ref name=mh33>Template:Harvnb</ref> The mane is raised during confrontations to make the aardwolf appear larger. It is missing the throat spot that others in the family have.<ref name=ingo/> Its lower leg (from the knee down) is all black, and its tail is bushy with a black tip.<ref name=w1/>

The aardwolf is about Template:Cvt long, excluding its bushy tail, which is about Template:Cvt long,<ref name=EB>Template:Harvnb</ref><ref name=w1/> and stands about Template:Cvt tall at the shoulders.<ref name="Colliers">Template:Harvnb</ref> An adult aardwolf weighs approximately Template:Cvt, sometimes reaching Template:Convert.<ref name=ingo>Template:Harvnb</ref> The aardwolves in the south of the continent tend to be slightly smaller (about Template:Cvt) than the eastern version (around Template:Cvt). This makes the aardwolf the smallest extant member of the Hyaenidae family.<ref name=mh33/> The front feet have five toes each, unlike the four-toed hyena.<ref name=EB/><ref>Template:Harvnb</ref> The skull is similar in shape to those of other hyenas, though much smaller,<ref name=Colliers/> and its cheek teeth are specialised for eating insects.<ref name=EB/> It still has canines, but unlike other hyenas, these teeth are used primarily for fighting and defense.<ref name=w1/> Its ears, which are large,<ref name=w1/> are very similar to those of the striped hyena.<ref name=ingo/>

As an aardwolf ages, it will typically lose some of its teeth, though this has little impact on its feeding habits due to the softness of the insects that it eats.<ref name="wff" />

Distribution and habitatEdit

Aardwolves live in open, dry plains and bushland, avoiding mountainous areas.<ref name=w1/> Due to their specific food requirements, they are found only in regions where termites of the family Hodotermitidae occur. Termites of this family depend on dead and withered grass and are most populous in heavily grazed grasslands and savannahs, including farmland. For most of the year, aardwolves spend time in shared territories consisting of up to a dozen dens, which are occupied for six weeks at a time.<ref name="wff" />

There are two distinct populations: one in Southern Africa, and another in East and Northeast Africa. The species does not occur in the intermediary miombo forests.

An adult pair, along with their most-recent offspring, occupies a territory of Template:Convert.<ref name=mh108>Template:Harvnb</ref>

Behavior and ecologyEdit

File:Aardwolf.jpg

Aardwolf at the San Antonio Zoo

Aardwolves are shy and nocturnal, sleeping in burrows by day.<ref name=EB/> They will, on occasion during the winter, become diurnal feeders. This happens during the coldest periods as they then stay in at night to conserve heat.<ref name=br30>Template:Harvnb</ref>

They are primarily solitary animals, though during mating season they form monogamous pairs which occupy a territory with their young.<ref name="Richardson">Richardson, P. R. K. "Aardwolf mating system: overt cuckoldry in an apparently monogamous mammal." South African Journal of Science 83.7 (1987): 405.</ref><ref name="koel">Template:Harvnb</ref> If their territory is infringed upon by another aardwolf, they will chase the intruder away for up to Template:Convert<ref name=PKR/> or to the border.<ref name=mh108/> If the intruder is caught, which rarely happens,<ref name=mh108/> a fight will occur, which is accompanied by soft clucking,<ref name=br31/> hoarse barking, and a type of roar.<ref name=PKR/> The majority of incursions occur during mating season, when they can occur once or twice per week.<ref name=PKR/> When food is scarce, the stringent territorial system may be abandoned and as many as three pairs may occupy a single territory.<ref name=PKR/>

The territory is marked by both sexes, as they both have developed anal glands from which they extrude a black substance that is smeared on rocks or grass stalks in Template:Convert-long streaks.<ref name=PKR/> Aardwolves also have scent glands on the forefoot and penile pad.<ref>Stoeckelhuber, Mechthild, Alexander Sliwa, and Ulrich Welsch. "Histo-physiology of the scent-marking glands of the penile pad, anal pouch, and the forefoot in the aardwolf (Proteles cristatus)." The anatomical record 259.3 (2000): 312-326.</ref> They often mark near termite mounds within their territory every 20 minutes or so. If they are patrolling their territorial boundaries, the marking frequency increases drastically, to once every Template:Convert. At this rate, an individual may mark 60 marks per hour,<ref name=PKR/> and upwards of 200 per night.<ref name=mh108/>

An aardwolf pair's territory may have up to 10 dens, and numerous middens where they dig small holes and bury their feces with sand.<ref>Template:Cite book</ref> Their dens are usually abandoned aardvark, springhare, or porcupine dens,<ref name=br31>Template:Harvnb</ref> or on occasion they are crevices in rocks. They will also dig their own dens, or enlarge dens started by springhares.<ref name=PKR/> They typically will only use one or two dens at a time, rotating through all of their dens every six months. During the summer, they may rest outside their den during the night and sleep underground during the heat of the day.

Aardwolves are not fast runners nor are they particularly adept at fighting off predators. Therefore, when threatened, the aardwolf may attempt to mislead its foe by doubling back on its tracks. If confronted, it may raise its mane in an attempt to appear more menacing. It also emits a foul-smelling liquid from its anal glands.<ref name=Colliers/>

FeedingEdit

The aardwolf feeds primarily on termites and more specifically on Trinervitermes.<ref name=mh71>Template:Harvnb</ref> This genus of termites has different species throughout the aardwolf's range. In East Africa, they eat Trinervitermes bettonianus, in central Africa, they eat Trinervitermes rhodesiensis, and in southern Africa, they eat T. trinervoides.<ref name=EB/><ref name=mh71/><ref name=PKR>Template:Harvnb</ref> Their technique consists of licking them off the ground as opposed to the aardvark, which digs into the mound.<ref name=br30/> They locate their food by sound and also from the scent secreted by the soldier termites.<ref name=PKR/> An aardwolf may consume up to 250,000 termites per night using its long, broad, sticky tongue.<ref name=mh71/><ref name="wff" />

They do not destroy the termite mound or consume the entire colony, thus ensuring that the termites can rebuild and provide a continuous supply of food. They often memorize the location of such nests and return to them every few months.<ref name=br31/> During certain seasonal events, such as the onset of the rainy season and the cold of midwinter, the primary termites become scarce, so the need for other foods becomes pronounced. During these times, the southern aardwolf will seek out Hodotermes mossambicus, a type of harvester termite<ref name=PKR/> active in the afternoon, which explains some of their diurnal behavior in the winter.<ref name=mh71/> The eastern aardwolf, during the rainy season, subsists on termites from the genera Odontotermes and Macrotermes.<ref name=mh71/> They are also known to feed on other insects and larvae, and, some sources mention, very occasionally small mammals and birds, but these constitute a very small percentage of their total diet.<ref name=PKR/> They use their wide tongues to lap surface foraging termites off of the ground and consume large quantities of sand in the process, which aids in digestion in the absence of teeth to break down their food.<ref name=Anderson1992>Template:Cite journal</ref>

Unlike other hyenas, aardwolves do not scavenge or kill larger animals.<ref name=w1/><ref name=br31/> Contrary to popular myths, aardwolves do not eat carrion, and if they are seen eating while hunched over a dead carcass, they are actually eating larvae and beetles.<ref name=w1/> Also, contrary to some sources, they do not like meat, unless it is finely ground or cooked for them.<ref name=w1/> The adult aardwolf was formerly assumed to forage in small groups,<ref name=Colliers/> but more recent research has shown that they are primarily solitary foragers,<ref name=koel/> necessary because of the scarcity of their insect prey. Their primary source, Trinervitermes, forages in small but dense patches of Template:Cvt.<ref name=PKR/> While foraging, the aardwolf can cover about Template:Cvt per hour, which translates to Template:Cvt per summer night and Template:Cvt per winter night.<ref name=w1/>

BreedingEdit

The breeding season varies depending on location, but normally takes place during autumn or spring. In South Africa, breeding occurs in early July.<ref name=mh108/> During the breeding season, unpaired male aardwolves search their own territory, as well as others, for a female to mate with. Dominant males also mate opportunistically with the females of less dominant neighboring aardwolves,<ref name=mh108/> which can result in conflict between rival males.<ref name=ingo/> Dominant males even go a step further and as the breeding season approaches, they make increasingly greater and greater incursions onto weaker males' territories. As the female comes into oestrus, they add pasting to their tricks inside of the other territories, sometimes doing so more in rivals' territories than their own.<ref name=mh108/> Females will also, when given the opportunity, mate with the dominant male, which increases the chances of the dominant male guarding "his" cubs with her.<ref name=mh108/> Copulation lasts between 1 and 4.5 hours.<ref name="Richardson"/><ref>Template:Cite thesis</ref>

Gestation lasts between 89 and 92 days,<ref name=ingo/><ref name=mh108/> producing two to five cubs (most often two or three) during the rainy season (October–December),<ref name=Colliers/> when termites are more active.<ref name=EB/> They are born with their eyes open, but initially are helpless,<ref name=PKR/> and weigh around Template:Cvt.<ref name=ingo/> The first six to eight weeks are spent in the den with their parents.<ref name=br31/> The male may spend up to six hours a night watching over the cubs while the mother is out looking for food.<ref name=mh108/><ref name=PKR/> After three months, they begin supervised foraging, and by four months are normally independent, though they often share a den with their mother until the next breeding season.<ref name=br31/> By the time the next set of cubs is born, the older cubs have moved on.<ref name=mh108/> Aardwolves generally achieve sexual maturity at one and a half to two years of age.<ref name=ingo/>

ConservationEdit

The aardwolf has not seen decreasing numbers and is relatively widespread throughout eastern Africa. They are not common throughout their range, as they maintain a density of no more than 1 per square kilometer, if food is abundant. Because of these factors, the IUCN has rated the aardwolf as least concern.<ref name=iucn /> In some areas, they are persecuted because of the mistaken belief that they prey on livestock; however, they are actually beneficial to the farmers because they eat termites that are detrimental.<ref name=PKR/> In other areas, the farmers have recognized this, but they are still killed, on occasion, for their fur. Dogs and insecticides<ref name=iucn /> are also common killers of the aardwolf.<ref name=br31/>