Red fox

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Red fox
Temporal range: 0.7 
Ma
Middle Pleistocene – present

Least Concern  (IUCN 3.1)[1]

Secure  (NatureServe)[2]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Vulpes
Species:
V. vulpes
Binomial name
Vulpes vulpes
Subspecies
Distribution of the red fox
  native
  introduced
  presence uncertain
Synonyms
  • Canis vulpes Linnaeus, 1758
  • Canis alopex Linnaeus, 1758

The red fox (Vulpes vulpes) is the largest of the

world's 100 worst invasive species".[4]

The red fox originated from smaller-sized ancestors from Eurasia during the Middle Villafranchian period,[5] and colonised North America shortly after the Wisconsin glaciation.[6] Among the true foxes, the red fox represents a more progressive form in the direction of carnivory.[7] Apart from its large size, the red fox is distinguished from other fox species by its ability to adapt quickly to new environments. Despite its name, the species often produces individuals with other colourings, including leucistic and melanistic individuals.[7] Forty-five subspecies are currently recognised,[8] which are divided into two categories: the large northern foxes and the small, basal southern grey desert foxes of Asia and North Africa.[7]

Red foxes are usually found in pairs or small groups consisting of families, such as a

Eurasian eagle owls,[11] and medium- and large-sized felids.[12]

The species has a long history of association with humans, having been extensively hunted as a pest and

colonised many suburban and urban areas. Domestication of the red fox is also underway in Russia, and has resulted in the domesticated silver fox
.

Terminology

Juvenile red foxes are known as kits

Males are called tods or dogs, females are called vixens, and young are known as cubs or kits.[14] Although the Arctic fox has a small native population in northern Scandinavia, and while the corsac fox's range extends into European Russia, the red fox is the only fox native to Western Europe, and so is simply called "the fox" in colloquial British English.

Etymology

The word "fox" comes from

Welsh name, llwynog, literally 'bushy', from llwyn 'bush'. Likewise, Portuguese: raposa from rabo 'tail', Lithuanian uodẽgis from uodegà 'tail', and Ojibwe waagosh from waa, which refers to the up and down "bounce" or flickering of an animal or its tail.[citation needed
]

The scientific term vulpes derives from the Latin word for fox, and gives the adjectives vulpine and vulpecular.[15]

Evolution

Comparative illustration of skulls of the red fox (left) and Rüppell's fox (right): note the more developed facial area of the former.

The red fox is considered to be a more specialised form of Vulpes than the

Afghan, corsac and Bengal fox species, in regards to their overall size and adaptation to carnivory; the skull displays far fewer neotenous traits than in other foxes, and its facial area is more developed.[7] It is, however, not as adapted for a purely carnivorous diet as the Tibetan fox.[7]

The sister lineage to the red fox is the Rüppell's fox, but the two species are surprisingly closely related through mitochondrial DNA markers, with Rüppell's fox nested inside the lineages of red foxes.[16][17] Such a nesting of one species within another is called paraphyly. Several hypotheses have been suggested to explain this,[16] including (1) recent divergence of Rüppell's fox from a red fox lineage, (2) incomplete lineage sorting, or introgression of mtDNA between the two species. Based on fossil record evidence, the last scenario seems most likely, which is further supported by the clear ecological and morphological differences between the two species.[citation needed]

Origins

The species is Eurasian in origin, and may have evolved from either Vulpes alopecoides or the related Chinese V. chikushanensis, both of which lived during the Middle

middens and refuse left by early human settlements. This has led to the theory that the red fox was hunted by primitive humans (as both a source of food and pelts); the possibility also exists of red foxes scavenging from middens or butchered animal carcasses.[21]

Colonisation of North America

Red foxes colonised the North American continent in two waves: before and during the

Sierra Nevada ranges, consisting of the smaller subspecies V. v. cascadensis, V. v. macroura, V. v. necator, and V. v. patwin. The latter clade has been separated from all other red fox populations since at least the last glacial maximum, and may possess unique ecological or physiological adaptations.[22]

Although European foxes (V. v. crucigera) were introduced to portions of the United States in the 1900s, recent genetic investigation indicates an absence of European fox mitochondrial haplotypes in any North American populations.

Monterey and north-coastal San Francisco Bay Area (including urban San Francisco and adjacent cities). In spite of the red fox's adaptability to city life, they are still found in somewhat greater numbers in the northern portions of California (north of the Bay Area) than in the south, as the wilderness is more alpine and isolated. The eastern red foxes appear to have mixed with the Sacramento Valley red fox (V. v. patwin) only in a narrow hybrid zone.[26] In addition, no evidence is seen of interbreeding of eastern American red foxes in California with the montane Sierra Nevada red fox (V. v. necator) or other populations in the Intermountain West (between the Rocky Mountains to the east and the Cascade and Sierra Nevada Mountains to the west).[27]

Subspecies

Skull of a northern fox
Skull of a southern grey desert fox

The 3rd edition of Mammal Species of the World[8] listed 45 subspecies as valid. In 2010, a distinct 46th subspecies, the Sacramento Valley red fox (V. v. patwin), which inhabits the grasslands of the Sacramento Valley, was identified through mitochondrial haplotype studies.[28] Castello (2018) recognized 30 subspecies of the Old World red fox and nine subspecies of the North American red fox as valid.[29]

Substantial

crossbred extensively with red foxes imported from Germany, France, Belgium, Sardinia and possibly Siberia and Scandinavia.[30]: 140  However, genetic studies suggest very little differences between red foxes sampled across Europe.[31][32] Lack of genetic diversity is consistent with the red fox being a highly agile species, with one red fox covering 320 km (200 mi) in under a year's time.[33]

Red fox subspecies in Eurasia and North Africa are divided into two categories:[7]

  • Northern foxes are large and brightly coloured.
  • Southern grey desert foxes include the Asian subspecies V. v. griffithi, V. v. pusilla, and V. v. flavescens. These foxes display transitional features between the northern foxes and other, smaller fox species; their skulls possess more primitive, neotenous traits than the northern foxes[7] and they are much smaller; the maximum sizes attained by southern grey desert foxes are invariably less than the average sizes of northern foxes. Their limbs are also longer and their ears larger.[7]

Red foxes living in Middle Asia show physical traits intermediate to the northern foxes and southern grey desert foxes.[7]

Description

Build

Red fox (left) and corsac fox (right) yawning

The red fox has an elongated body and relatively short limbs. The tail, which is longer than half the body length

testes of males are smaller than those of Arctic foxes.[7]

Their

canine teeth are relatively long. Sexual dimorphism of the skull is more pronounced than in corsac foxes, with female red foxes tending to have smaller skulls than males, with wider nasal regions and hard palates, as well as having larger canines.[7] Their skulls are distinguished from those of dogs by their narrower muzzles, less crowded premolars, more slender canine teeth, and concave rather than convex profiles.[9]

Dimensions

Red foxes are the largest species of the genus Vulpes.[46] However, relative to dimensions, red foxes are much lighter than similarly sized dogs of the genus Canis. Their limb bones, for example, weigh 30 percent less per unit area of bone than expected for similarly sized dogs.[47] They display significant individual, sexual, age and geographical variation in size. On average, adults measure 35–50 cm (14–20 in) high at the shoulder and 45–90 cm (18–35 in) in body length with tails measuring 30–55.5 cm (11.8–21.9 in). The ears measure 7.7–12.5 cm (3.0–4.9 in) and the hind feet 12–18.5 cm (4.7–7.3 in). Weights range from 2.2–14 kg (4.9–30.9 lb), with vixens typically weighing 15–20% less than males.[48][49] Adult red foxes have skulls measuring 129–167 mm (5.1–6.6 in), while those of vixens measure 128–159 mm (5.0–6.3 in).[7] The forefoot print measures 60 mm (2.4 in) in length and 45 mm (1.8 in) in width, while the hind foot print measures 55 mm (2.2 in) long and 38 mm (1.5 in) wide. They trot at a speed of 6–13 km/h (3.7–8.1 mph), and have a maximum running speed of 50 km/h (31 mph). They have a stride of 25–35 cm (9.8–13.8 in) when walking at a normal pace.[47]: 36  North American red foxes are generally lightly built, with comparatively long bodies for their mass and have a high degree of sexual dimorphism. British red foxes are heavily built, but short, while continental European red foxes are closer to the general average among red fox populations.[50] The largest red fox on record in Great Britain was a 1.4 m (4 ft 7 in) long male, that weighed 17.2 kg (38 lb), killed in Aberdeenshire, Scotland, in early 2012.[51]

Fur

A red fox in its winter coat in the Rocky Mountains of Colorado U.S.A.

The winter fur is dense, soft, silky and relatively long. For the northern foxes, the fur is very long, dense and fluffy, but it is shorter, sparser and coarser in southern forms.

supracaudal gland, is usually present at the base of the tail. The tip of the tail is white.[7]

Colour morphs

Various red fox colour morphs

Atypical colouration in the red fox usually represents stages toward full melanism,[7] and mostly occurs in cold regions.[10]

Colour morph Image Description
Red The typical colouration (see fur above)
Smokey The rump and spine is brown or grey with light yellowish bands on the guard hairs. The cross on the shoulders is brown, rusty brown or reddish-brown. The limbs are brown.[7]
Cross
The fur has a darker colouration than the colour morph listed directly above. The rump and lower back are dark brown or dark grey, with varying degrees of silver on the guard hairs. The cross on the shoulders is black or brown, sometimes with light silvery fur. The head and feet are brown.[7]
Blackish-brown The melanistic colour morph of the Eurasian red foxes. Has blackish-brown or black skin with a light brownish tint. The skin area usually has a variable admixture of silver. Reddish hairs are either completely absent or in small quantities.[7]
Silver The melanistic colour morph of the North American red foxes, but introduced to the Old World by the fur trade. Characterised by pure black colour with skin that usually has a variable admixture of silver (covering 25–100% of the skin area)[7]
Platinum Distinguished from the silver colour morph by its pale, almost silvery-white fur with a bluish cast[13]: 251 
Amber
Samson Distinguished by its woolly pelt, which lacks guard hairs[13]: 230 

Senses

Red foxes have

auditory perception is acute, being able to hear black grouse changing roosts at 600 paces, the flight of crows at 0.25–0.5 km (0.16–0.31 mi) and the squeaking of mice at about 100 m (330 ft).[7] They are capable of locating sounds to within one degree at 700–3,000 Hz, though less accurately at higher frequencies.[45] Their sense of smell is good, but weaker than that of specialised dogs.[7]

Scent glands

Red foxes have a pair of

violets.[7] The presence of foot glands is equivocal. The interdigital cavities are deep, with a reddish tinge and smell strongly. Sebaceous glands are present on the angle of the jaw and mandible.[9]

Distribution and habitat

Multi-coloured North American red fox

The red fox is a wide-ranging species. Its range covers nearly 70,000,000 km2 (27,000,000 sq mi) including as far north as the Arctic Circle. It occurs all across Europe, in Africa north of the Sahara Desert, throughout Asia apart from extreme Southeast Asia, and across North America apart from most of the southwestern United States and Mexico. It is absent in Greenland, Iceland, the Arctic islands, the most northern parts of central Siberia, and in extreme deserts.[1] It is not present in New Zealand and is classed as a "prohibited new organism" under the Hazardous Substances and New Organisms Act 1996, which does not allow import.[55]

Australia

In Australia, estimates in 2012 indicated that there were more than 7.2 million red foxes,

niche differentiation with both the feral dog and the feral cat. Consequently, the fox has become one of the continent's most destructive invasive species.[citation needed
]

The red fox has been implicated in the extinction or decline of several native Australian species, particularly those of the family

Sardinia, Italy

The origin of the ichnusae subspecies in Sardinia, Italy is uncertain, as it is absent from Pleistocene deposits in their current homeland. It is possible it originated during the Neolithic following its introduction to the island by humans. It is likely then that Sardinian fox populations stem from repeated introductions of animals from different localities in the Mediterranean. This latter theory may explain the subspecies' phenotypic diversity.[21]

Behaviour

Social and territorial behaviour

Red fox pressed against the trunk of a pine tree in Ilmatsalu, Estonia
Red foxes grooming each other
A pair of European red foxes (V. v. crucigera) at the British Wildlife Centre, Surrey, England
A red fox marking its territory

Red foxes either establish stable home ranges within particular areas or are itinerant with no fixed abode.

dominant animals, but are mostly young from the previous year, who act as helpers in rearing the breeding vixen's kits. Alternatively, their presence has been explained as being in response to temporary surpluses of food unrelated to assisting reproductive success. Non-breeding vixens will guard, play, groom, provision and retrieve kits,[9] an example of kin selection. Red foxes may leave their families once they reach adulthood if the chances of winning a territory of their own are high. If not, they will stay with their parents, at the cost of postponing their own reproduction.[47]
: 140–141 

Reproduction and development

Red fox kits
Red foxes mating

Red foxes reproduce once a year in spring. Two months prior to

gestation period lasts 49–58 days.[7] Though foxes are largely monogamous,[69] DNA evidence from one population indicated large levels of polygyny, incest and mixed paternity litters.[9] Subordinate vixens may become pregnant, but usually fail to whelp, or have their kits killed postpartum by either the dominant female or other subordinates.[9]

Red fox kits coming out of their den

The average litter size consists of four to six kits, though litters of up to 13 kits have occurred.[7] Large litters are typical in areas where fox mortality is high.[47]: 93  Kits are born blind, deaf and toothless, with dark brown fluffy fur. At birth, they weigh 56–110 g (2.0–3.9 oz) and measure 14.5 cm (5.7 in) in body length and 7.5 cm (3.0 in) in tail length. At birth, they are short-legged, large-headed and have broad chests.[7] Mothers remain with the kits for 2–3 weeks, as they are unable to thermoregulate. During this period, the fathers or barren vixens feed the mothers.[9] Vixens are very protective of their kits, and have been known to even fight off terriers in their defence.[30]: 21–22  If the mother dies before the kits are independent, the father takes over as their provider.[30]: 13  The kits' eyes open after 13–15 days, during which time their ear canals open and their upper teeth erupt, with the lower teeth emerging 3–4 days later.[7] Their eyes are initially blue, but change to amber at 4–5 weeks. Coat colour begins to change at three weeks of age, when the black eye streak appears. By one month, red and white patches are apparent on their faces. During this time, their ears erect and their muzzles elongate.[9] Kits begin to leave their dens and experiment with solid food brought by their parents at the age of 3–4 weeks. The lactation period lasts 6–7 weeks.[7] Their woolly coats begin to be coated by shiny guard hairs after 8 weeks.[9] By the age of 3–4 months, the kits are long-legged, narrow-chested and sinewy. They reach adult proportions at the age of 6–7 months.[7] Some vixens may reach sexual maturity at the age of 9–10 months, thus bearing their first litters at one year of age.[7] In captivity, their longevity can be as long as 15 years, though in the wild they typically do not survive past 5 years of age.[70]

Denning behaviour

Side and above view of a red fox den

Outside the

latrines, red foxes habitually leave pieces of prey around their dens.[30]: 15–17  The average sleep time of a captive red fox is 9.8 hours per day.[71]

Communication

Body language

A European fox (V. v. crucigera) in an inquisitive posture

Red fox body language consists of movements of the ears, tail and postures, with their body markings emphasising certain gestures. Postures can be divided into aggressive/dominant and fearful/submissive categories. Some postures may blend the two together.[47]: 42–43  Inquisitive foxes will rotate and flick their ears whilst sniffing. Playful individuals will perk their ears and rise on their hind legs. Male foxes courting females, or after successfully evicting intruders, will turn their ears outwardly, and raise their tails in a horizontal position, with the tips raised upward. When afraid, red foxes grin in submission, arching their backs, curving their bodies, crouching their legs and lashing their tails back and forth with their ears pointing backwards and pressed against their skulls. When merely expressing submission to a dominant animal, the posture is similar, but without arching the back or curving the body. Submissive foxes will approach dominant animals in a low posture, so that their muzzles reach up in greeting. When two evenly matched foxes confront each other over food, they approach each other sideways and push against each other's flanks, betraying a mixture of fear and aggression through lashing tails and arched backs without crouching and pulling their ears back without flattening them against their skulls. When launching an assertive attack, red foxes approach directly rather than sideways, with their tails aloft and their ears rotated sideways.[47] During such fights, red foxes will stand on each other's upper bodies with their forelegs, using open mouthed threats. Such fights typically only occur among juveniles or adults of the same sex.[9]

Vocalisations

A pair of Wasatch Mountains foxes (V. v. macroura) squabbling

Red foxes have a wide vocal range, and produce different sounds spanning five octaves, which grade into each other.[47]: 28  Recent analyses identify 12 different sounds produced by adults and 8 by kits.[9] The majority of sounds can be divided into "contact" and "interaction" calls. The former vary according to the distance between individuals, while the latter vary according to the level of aggression.[47]: 28 

  • Contact calls: The most commonly heard contact call is a three to five syllable barking "wow wow wow" sound, which is often made by two foxes approaching one another. This call is most frequently heard from December to February (when they can be confused with the territorial calls of tawny owls). The "wow wow wow" call varies according to individual; captive foxes have been recorded to answer pre-recorded calls of their pen-mates, but not those of strangers. Kits begin emitting the "wow wow wow" call at the age of 19 days, when craving attention. When red foxes draw close together, they emit trisyllabic greeting warbles similar to the clucking of chickens. Adults greet their kits with gruff huffing noises.[47]: 28 
  • Interaction calls: When greeting one another, red foxes emit high pitched whines, particularly submissive animals. A submissive fox approached by a dominant animal will emit a ululating siren-like shriek. During aggressive encounters with conspecifics, they emit a throaty rattling sound, similar to a ratchet, called "gekkering". Gekkering occurs mostly during the courting season from rival males or vixens rejecting advances.[47]: 28 

Another call that does not fit into the two categories is a long, drawn-out, monosyllabic "waaaaah" sound. As it is commonly heard during the breeding season, it is thought to be emitted by vixens summoning males. When danger is detected, foxes emit a monosyllabic bark. At close quarters, it is a muffled cough, while at long distances it is sharper. Kits make warbling whimpers when nursing, these calls being especially loud when they are dissatisfied.[47]: 28 

Ecology

Diet, hunting and feeding behaviour

coypu

Red foxes are

grasses, sedges and tubers.[10]

Red foxes are implicated in the predation of

song birds, hares, rabbits, muskrats and young ungulates, particularly in preserves, reserves and hunting farms where ground-nesting birds are protected and raised, as well as in poultry farms.[7]

While the popular consensus is that

olfaction is very important for hunting,[75] two studies that experimentally investigated the role of olfactory, auditory and visual cues found that visual cues are the most important ones for hunting in red foxes[76] and coyotes.[77][78]

Red foxes prefer to hunt in the early morning hours before sunrise and late evening.[7] Although they typically forage alone, they may aggregate in resource-rich environments.[70] When hunting mouse-like prey, they first pinpoint their prey's location by sound, then leap, sailing high above their quarry, steering in mid-air with their tails, before landing on target up to 5 m (16 ft) away.[1] They typically only feed on carrion in the late evening hours and at night.[7] They are extremely possessive of their food and will defend their catches from even dominant animals.[47]: 58  Red foxes may occasionally commit acts of surplus killing; during one breeding season, four red foxes were recorded to have killed around 200 black-headed gulls each, with peaks during dark, windy hours when flying conditions were unfavourable. Losses to poultry and penned game birds can be substantial because of this.[9][47]: 164  Red foxes seem to dislike the taste of moles, but will nonetheless catch them alive and present them to their kits as playthings.[47]: 41 

A 2008–2010 study of 84 red foxes in the Czech Republic and Germany found that successful hunting in long vegetation or under snow appeared to involve an alignment of the red fox with the Earth's magnetic field.[79][80]

Enemies and competitors

A red fox confronting a grey fox

Red foxes typically dominate other fox species. Arctic foxes generally escape competition from red foxes by living farther north, where food is too scarce to support the larger-bodied red species. Although the red species' northern limit is linked to the availability of food, the Arctic species' southern range is limited by the presence of the former. Red and Arctic foxes were both introduced to almost every island from the Aleutian Islands to the Alexander Archipelago during the 1830s–1930s by fur companies. The red foxes invariably displaced the Arctic foxes, with one male red fox having been reported to have killed off all resident Arctic foxes on a small island in 1866.[47] Where they are sympatric, Arctic foxes may also escape competition by feeding on lemmings and flotsam rather than voles, as favoured by red foxes. Both species will kill each other's kits, given the opportunity.[7] Red foxes are serious competitors of corsac foxes, as they hunt the same prey all year. The red species is also stronger, is better adapted to hunting in snow deeper than 10 cm (3.9 in) and is more effective in hunting and catching medium-sized to large rodents. Corsac foxes seem to only outcompete red foxes in semi-desert and steppe areas.[7][81] In Israel, Blanford's foxes escape competition with red foxes by restricting themselves to rocky cliffs and actively avoiding the open plains inhabited by red foxes.[47]: 84–85  Red foxes dominate kit and swift foxes. Kit foxes usually avoid competition with their larger cousins by living in more arid environments, though red foxes have been increasing in ranges formerly occupied by kit foxes due to human-induced environmental changes. Red foxes will kill both species and compete with them for food and den sites.[10] Grey foxes are exceptional, as they dominate red foxes wherever their ranges meet. Historically, interactions between the two species were rare, as grey foxes favoured heavily wooded or semiarid habitats as opposed to the open and mesic ones preferred by red foxes. However, interactions have become more frequent due to deforestation, allowing red foxes to colonise grey fox-inhabited areas.[10]

Female red fox and kits interacting with a male golden jackal in south-western Germany

Wolves may kill and eat red foxes in disputes over carcasses.[7][82] In areas in North America where red fox and coyote populations are sympatric, red fox ranges tend to be located outside coyote territories. The principal cause of this separation is believed to be active avoidance of coyotes by the red foxes. Interactions between the two species vary in nature, ranging from active antagonism to indifference. The majority of aggressive encounters are initiated by coyotes, and there are few reports of red foxes acting aggressively toward coyotes except when attacked or when their kits were approached. Foxes and coyotes have sometimes been seen feeding together.[83] In Israel, red foxes share their habitat with golden jackals. Where their ranges meet, the two canids compete due to near-identical diets. Red foxes ignore golden jackal scents or tracks in their territories and avoid close physical proximity with golden jackals themselves. In areas where golden jackals become very abundant, the population of red foxes decreases significantly, apparently because of competitive exclusion.[84] There is however one record of multiple red foxes interacting peacefully with a golden jackal in southwestern Germany.[85]

A golden eagle feeding on a red fox

Red foxes dominate

raccoon dogs, sometimes killing their kits or biting adults to death. Cases are known of red foxes killing raccoon dogs after entering their dens. Both species compete for mouse-like prey. This competition reaches a peak during early spring when food is scarce. In Tatarstan, red fox predation accounted for 11.1% of deaths among 54 raccoon dogs and amounted to 14.3% of 186 raccoon dog deaths in northwestern Russia.[7]

Red foxes may kill small

Eurasian badgers may live alongside red foxes in isolated sections of large burrows.[7] It is possible that the two species tolerate each other out of mutualism; red foxes provide Eurasian badgers with food scraps, while Eurasian badgers maintain the shared burrow's cleanliness.[30]: 15  However, cases are known of Eurasian badgers driving vixens from their dens and destroying their litters without eating them.[87] Wolverines may kill red foxes, often while the latter is sleeping or near carrion.[7]: 546  Red foxes, in turn, may kill young wolverines.[88]

Red foxes may compete with striped hyenas on large carcasses. Red foxes may give way to striped hyenas on unopened carcasses, as the latter's stronger jaws can easily tear open flesh that is too tough for red foxes. Red foxes may harass striped hyenas, using their smaller size and greater speed to avoid the hyena's attacks. Sometimes, red foxes seem to deliberately torment striped hyenas even when there is no food at stake. Some red foxes may mis-time their attacks and are killed.[47]: 77–79  Red fox remains are often found in striped hyena dens and striped hyenas may steal red foxes from traps.[7]

In Eurasia, red foxes may be preyed upon by leopards, caracals and Eurasian lynxes. The Eurasian lynxes chase red foxes into deep snow, where their long legs and larger paws give them an advantage over red foxes, especially when the depth of the snow exceeds one meter.[7] In the Velikoluksky District in Russia, red foxes are absent or are seen only occasionally where Eurasian lynxes establish permanent territories.[7] Researchers consider Eurasian lynxes to represent considerably less danger to red foxes than wolves do.[7] North American felid predators of red foxes include cougars, Canada lynxes and bobcats.[44]

Red foxes compete with various birds of prey such as common buzzards (Buteo buteo) and northern goshawks (Accipiter gentilis) and even steal their kills.[89][90] In turn, golden eagles (Aquila chrysaetos) regularly takes young red foxes and prey on adults if needed.[91][92] Other large eagles such as wedge-tailed eagles (Aquila audax), eastern imperial eagles (Aquila heliaca), white-tailed eagles (Haliaeetus albicilla), and steller's sea eagles (Haliaeetus pelagicus) have also been known to kill red foxes less frequently.[93][94][95][96][97] Additionally, large owls such as Eurasian eagle-owls (Bubo bubo) and snowy owls (Bubo scandiacus) will prey on young foxes, and adults on exceptional occasions.[98][99][100]

Diseases and parasites

A European fox (V. v. crucigera) with mange

Red foxes are the most important

vectors in spreading erysipelas, brucellosis and tick-borne encephalitis. A mysterious fatal disease near Lake Sartlan in the Novosibirsk Oblast was noted among local red foxes, but the cause was undetermined. The possibility was considered that it was caused by an acute form of encephalomyelitis, which was first observed in captive-bred silver foxes. Individual cases of foxes infected with Yersinia pestis are known.[7]

Red foxes are not readily prone to infestation with

Up to 60

Pachysentis ehrenbergi in both in Egypt.[105]

Relationships with humans

In folklore, religion and mythology

Reynard the Fox in an 1869 children's book

Red foxes feature prominently in the folklore and mythology of human cultures with which they are sympatric. In Greek mythology, the Teumessian fox,[106] or Cadmean vixen, was a gigantic fox that was destined never to be caught. The fox was one of the children of Echidna.[107]

In

witches were thought to take the shape of foxes to steal butter from their neighbours.[108] In later European folklore, the figure of Reynard the Fox symbolises trickery and deceit. He originally appeared (then under the name of "Reinardus") as a secondary character in the 1150 poem "Ysengrimus". He reappeared in 1175 in Pierre Saint Cloud's Le Roman de Renart, and made his debut in England in Geoffrey Chaucer's The Nun's Priest's Tale. Many of Reynard's adventures may stem from actual observations on fox behaviour; he is an enemy of the wolf and has a fondness for blackberries and grapes.[47]
: 32–33 

Chinese folk tales tell of fox-spirits called

Arab folklore, the fox is considered a cowardly, weak, deceitful, and cunning animal, said to feign death by filling its abdomen with air to appear bloated, then lies on its side, awaiting the approach of unwitting prey.[41] The animal's cunning was noted by the authors of the Bible who applied the word "fox" to false prophets (Ezekiel 13:4) and the hypocrisy of Herod Antipas (Luke 13:32).[111]

The cunning Fox is commonly found in

Yurok tribe believed that Fox, in anger, captured the Sun, and tied him to a hill, causing him to burn a great hole in the ground. An Inuit story tells of how Fox, portrayed as a beautiful woman, tricks a hunter into marrying her, only to resume her true form and leave after he offends her. A Menominee story tells of how Fox is an untrustworthy friend to Wolf.[112]

Hunting

Beagle and Fox (1885) by Bruno Liljefors

The earliest historical records of fox hunting come from the 4th century BC;

Edward, Second Duke of York assisted the climb of foxes as more prestigious quarries in his The Master of Game. By the Renaissance, fox hunting became a traditional sport of the nobility. After the English Civil War caused a drop in deer populations, fox hunting grew in popularity. By the mid-1600s, Great Britain was divided into fox hunting territories, with the first fox hunting clubs being formed (the first was the Charlton Hunt Club in 1737). The popularity of fox hunting in Great Britain reached a peak during the 1700s.[47]: 21  Although already native to North America, red foxes from England were imported for sporting purposes to Virginia and Maryland in 1730 by prosperous tobacco planters.[113] These American fox hunters considered the red fox more sporting than the grey fox.[113]

The grays furnished more fun, the reds more excitement. The grays did not run so far, but usually kept near home, going in a circuit of six or eight miles. 'An old red, generally so called irrespective of age, as a tribute to his prowess, might lead the dogs all day, and end by losing them as evening fell, after taking them a dead stretch for thirty miles. The capture of a gray was what men boasted of; a chase after 'an old red' was what they 'yarned' about.[113]

Red foxes are still widely persecuted as pests, with human-caused deaths among the highest causes of mortality in the species. Annual red fox kills are: UK 21,500–25,000 (2000); Germany 600,000 (2000–2001); Austria 58,000 (2000–2001); Sweden 58,000 (1999–2000); Finland 56,000 (2000–2001); Denmark 50,000 (1976–1977); Switzerland 34,832 (2001); Norway 17,000 (2000–2001); Saskatchewan (Canada) 2,000 (2000–2001); Nova Scotia (Canada) 491 (2000–2001); Minnesota (US) 4,000–8,000 (average annual trapping harvest 2002–2009);[114] New Mexico (US) 69 (1999–2000).[86]

Fur use

Red fox pelts
A red fox in a fur farm in Vörå, Finland

Red foxes are among the most important

coats. They are principally used as trimming for both cloth coats and fur garments, including evening wraps.[13]: 229–230  The pelts of silver foxes are popular as capes,[13]: 246  while cross foxes are mostly used for scarves and rarely for trimming.[13]: 252  The number of sold fox scarves exceeds the total number of scarves made from other fur-bearers. However, this amount is overshadowed by the total number of red fox pelts used for trimming purposes.[13]: 229–230  The silver colour morphs are the most valued by furriers, followed by the cross colour morphs and the red colour morphs, respectively.[30]: 207  In the early 1900s, over 1,000 American red fox skins were imported to Great Britain annually, while 500,000 were exported annually from Germany and Russia.[30]: 6  The total worldwide trade of wild red foxes in 1985–86 was 1,543,995 pelts. Red foxes amounted to 45% of U.S. wild-caught pelts worth $50 million.[86] Pelt prices are increasing, with 2012 North American wholesale auction prices averaging $39 and 2013 prices averaging $65.78.[115]

North American red foxes, particularly those of northern Alaska, are the most valued for their fur, as they have guard hairs of a silky texture which, after dressing, allow the wearer unrestricted mobility. Red foxes living in southern Alaska's coastal areas and the Aleutian Islands are an exception, as they have extremely coarse pelts that rarely exceed one-third of the price of their northern Alaskan cousins.[13]: 231  Most European peltries have coarse-textured fur compared to North American varieties. The only exceptions are the Nordic and Far Eastern Russian peltries, but they are still inferior to North American peltries in terms of silkiness.[13]: 235 

Livestock and pet predation

A carcass of a lamb near a red fox den
A red fox in a Birmingham garden investigating a rabbit hutch

Red foxes may on occasion prey on lambs. Usually, lambs targeted by red foxes tend to be physically weakened specimens, but not always. Lambs belonging to small breeds, such as the Scottish Blackface, are more vulnerable than larger breeds, such as the Merino. Twins may be more vulnerable to red foxes than singlets, as ewes cannot effectively defend both simultaneously. Crossbreeding small, upland ewes with larger, lowland rams can cause difficult and prolonged labour for ewes due to the heaviness of the resulting offspring, thus making the lambs more at risk to red fox predation. Lambs born from gimmers (ewes breeding for the first time) are more often killed by red foxes than those of experienced mothers, who stick closer to their young.[47]: 166–167 

Red foxes may prey on domestic rabbits and guinea pigs if they are kept in open runs or are allowed to range freely in gardens. This problem is usually averted by housing them in robust hutches and runs. Urban red foxes frequently encounter cats and may feed alongside them. In physical confrontations, the cats usually have the upper hand. Authenticated cases of red foxes killing cats usually involve kittens. Although most red foxes do not prey on cats, some may do so and may treat them more as competitors rather than food.[47]: 180–181 

Taming and domestication

A young boy holding a tame red fox kit

In their unmodified wild state, red foxes are generally unsuitable as pets.[116] Many supposedly abandoned kits are adopted by well-meaning people during the spring period, though it is unlikely that vixens would abandon their young. Actual orphans are rare and the ones that are adopted are likely kits that simply strayed from their den sites.[117] Kits require almost constant supervision; when still suckling, they require milk at four-hour intervals day and night. Once weaned, they may become destructive to leather objects, furniture and electric cables.[47]: 56  Though generally friendly toward people when young, captive red foxes become fearful of humans, save for their handlers, once they reach 10 weeks of age.[47]: 61  They maintain their wild counterparts' strong instinct of concealment and may pose a threat to domestic birds, even when well-fed.[30]: 122  Although suspicious of strangers, they can form bonds with cats and dogs, even ones bred for fox hunting. Tame red foxes were once used to draw ducks close to hunting blinds.[30]: 132–133 

White to black individual red foxes have been selected and raised on fur farms as "

Russian geneticist Dmitry Belyayev who, over a 40-year period, bred several generations selecting only those individuals that showed the least fear of humans. Eventually, Belyayev's team selected only those that showed the most positive response to humans, thus resulting in a population of silver foxes whose behaviour and appearance was significantly changed. After about 10 generations of controlled breeding, these foxes no longer showed any fear of humans and often wagged their tails and licked their human caretakers to show affection. These behavioural changes were accompanied by physical alterations, which included piebald coats, floppy ears in kits and curled tails, similar to the traits that distinguish domestic dogs from grey wolves.[118]

Urban red foxes

Distribution

Red foxes have been exceedingly successful in colonising built-up environments, especially lower-density suburbs,

Zurich, Switzerland, they only started appearing in the 1980s.[119] Urban red foxes are most common in residential suburbs consisting of privately owned, low-density housing. They are rare in areas where industry, commerce or council-rented houses predominate.[45] In these latter areas, the distribution is of a lower average density because they rely less on human resources; the home range of these foxes average from 80–90 ha (0.80–0.90 km2; 200–220 acres), whereas those in more residential areas average from 25–40 ha (0.25–0.40 km2; 62–99 acres).[120]

In 2006, it was estimated that there were 10,000 red foxes in London.[121] City-dwelling red foxes may have the potential to consistently grow larger than their rural counterparts as a result of abundant scraps and a relative lack of predators. In cities, red foxes may scavenge food from litter bins and bin bags, although much of their diet is similar to rural red foxes.[citation needed]

Behaviour

Urban red foxes are most active at dusk and dawn, doing most of their hunting and scavenging at these times. It is uncommon to spot them during the day, but they can be caught sunbathing on roofs of houses or sheds. Urban red foxes will often make their homes in hidden and undisturbed spots in urban areas as well as on the edges of a city, visiting at night for sustenance. They sleep at night in dens. While urban red foxes will scavenge successfully in the city (and the red foxes tend to eat anything that humans eat) some urban residents will deliberately leave food out for the animals, finding them endearing. Doing this regularly can attract urban red foxes to one's home; they can become accustomed to human presence, warming up to their providers by allowing themselves to be approached and in some cases even played with, particularly young kits.[120]

Urban red fox control

Urban red foxes can cause problems for local residents. They have been known to steal chickens, disrupt rubbish bins and damage gardens. Most complaints about urban red foxes made to local authorities occur during the breeding season in late January/early February or from late April to August when the new kits are developing.[120] In the U.K., hunting red foxes in urban areas is banned and shooting them in an urban environment is not suitable. One alternative to hunting urban red foxes has been to trap them, which appears to be a more viable method.[122] However, killing red foxes has little effect on the population in an urban area; those that are killed are very soon replaced, either by new kits during the breeding season or by other red foxes moving into the territory of those that were killed. A more effective method of urban red fox control is to deter them from the specific areas they inhabit. Deterrents such as creosote, diesel oil, or ammonia can be used. Cleaning up and blocking access to den locations can also discourage an urban red fox's return.[120]

Relationship between urban and rural red foxes

In January 2014 it was reported that "Fleet", a relatively tame urban red fox tracked as part of a wider study by the

GPS collar stopped transmitting due to suspected water damage. Along with setting a record for the longest journey undertaken by a tracked red fox in the United Kingdom, his travels have highlighted the fluidity of movement between rural and urban red fox populations.[123][124]

  • An urban red fox crossing a street
    An urban red fox crossing a street
  • An urban red fox in central London
    An urban red fox in central London
  • An urban red fox eating from a bag of biscuits in Dorset, England
    An urban red fox eating from a bag of biscuits in Dorset, England
  • "Fleet", the urban red fox from the BBC TV series Winterwatch
    "Fleet", the urban red fox from the BBC TV series Winterwatch
  • An urban red fox with a discarded KFC bag

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Further reading

External links