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Red fox

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From Wikipedia, the free encyclopedia
Species of mammal
For other uses, seeRed Fox (disambiguation).

Red fox
Temporal range:0.7 MaMiddlePleistocene – present
in Rautas fjällurskog nature reserve, Sweden
Secure
Secure (NatureServe)[2]
Scientific classificationEdit this classification
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 vulpesLinnaeus, 1758
  • Canis alopexLinnaeus, 1758

Thered fox (Vulpes vulpes) is the largest of thetrue foxes and one of the most widely distributed members of the orderCarnivora. It is present across the entireNorthern Hemisphere, including most ofNorth America,Europe andAsia, as well as parts ofNorth Africa. Its range has expanded alongside human settlement, with the species having beenintroduced to Australia, where it preys on native small and medium-sized rodents andmarsupials. The red fox is listed as ofleast concern on theIUCN Red List.[1] Because of its impact on native species, it is also included on the list of the "world's 100 worst invasive species".[4]

The red fox originated in Eurasia during theMiddle Pleistocene at least 400,000 years ago[5] and later colonised North America sometime prior to 130,000 years ago.[6] Among the true foxes, the red fox represents a more progressive form in the direction ofcarnivory.[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, includingleucistic andmelanistic individuals.[7] Forty-fivesubspecies 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 in small family groups consisting of amated pair and their young, or a male with several females related to each other. The young of the mated pair stay with their parents to help care for the new kits.[9] The species primarily feeds on small rodents, though it may also targetrabbits,squirrels,game birds,reptiles,invertebrates[7] and youngungulates.[7] Fruit and vegetable matter is also eaten sometimes.[10] Although the red fox tends to kill smaller predators, including other fox species, it is vulnerable to attack from larger predators, such aswolves,coyotes,golden jackals, large predatory birds such asgolden eagles andEurasian eagle-owls,[11] and medium- and large-sizedfelids.[12]

The species has a long history of association with humans. It has been extensively hunted as a pest andfurbearer for many centuries, and is represented in human folklore and mythology. Due to its widespread distribution and large population, the red fox is one of the most important fur-bearing animals harvested for thefur trade.[13]: 229–230  Too small to pose a significant threat to humans, it has greatly benefited from the presence of human habitation, and has successfullycolonised manysuburban andurban areas. The domestication of the red fox is also underway inRussia, and has resulted in thedomesticated silver fox.

Terminology

[edit]
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 theArctic fox has a small native population in northern Scandinavia, and while thecorsac fox's range extends intoEuropean Russia, the red fox is the only fox native to Western Europe. For this reason, it is simply called "the fox" in colloquial British English.

Etymology

[edit]

The word "fox" comes fromOld English, which in turn derived fromProto-Germanic *fuhsaz. Compare withWest Frisianfoks,Dutchvos, andGermanFuchs. This, in turn, derives fromProto-Indo-European *puḱ- 'thick-haired; tail'. Compare to theHindipū̃ch 'tail',Tocharian Bpäkā 'tail; chowrie', andLithuanianpūkas 'fur / fluff'. The fox's bushy tail also forms the basis for itsWelsh name,llwynog, literally 'bushy', fromllwyn 'bush'. Likewise,Portuguese:raposa fromrabo 'tail', Lithuanianuodẽgis fromuodegà 'tail', andOjibwewaagosh fromwaa, which refers to the up and down "bounce" or flickering of an animal or its tail.[citation needed]

The scientific termvulpes derives from the Latin word for fox, and gives the adjectivesvulpine andvulpecular.[15]

Evolutionary history

[edit]
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 ofVulpes than theAfghan,corsac andBengal fox species in terms of their overall size and adaptation tocarnivory; the skull displays far fewerneotenous traits than those of other foxes, and the facial area is more developed.[7] However, it is not as well adapted to a purely carnivorous diet as theTibetan fox.[7]

The sister lineage to the red fox is theRüppell's fox, but the two species are surprisingly closely related throughmitochondrial DNA markers, with Rüppell's fox nested within the red fox lineage.[16][17] Such a nesting of one species within another is calledparaphyly. 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, orintrogression of mtDNA between the two species. Based on the fossil record, the last scenario seems the most likely. This is further supported by the clear ecological and morphological differences between the two species.[citation needed]

Origins

[edit]

The species is Eurasian in origin, and may have evolved from eitherVulpes alopecoides or the related ChineseV. chikushanensis, both of which lived during the MiddleVillafranchian of thePleistocene Epoch.[19] The earliest fossil specimens ofV. vulpes were uncovered inBaranya County,Hungary, dating from 3.4 to 1.8 million years ago.[20] The ancestral red fox was likely more diminutive compared to today's extant foxes, as the earliest red fox fossils have shown a smaller build than living specimens.[19]: 115–116  The earliest fossil remains of the modern species date back to the mid-Pleistocene,[21] found in association withmiddens 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.[22]

Colonisation of North America

[edit]

Red foxes colonised the North American continent in two waves: before and during theIllinoian glaciation, and during theWisconsinan glaciation.[23]Gene mapping demonstrates that red foxes in North America have been isolated from their Old World counterparts for over 400,000 years, thus raising the possibility thatspeciation has occurred, and that the previousbinomial name ofVulpes fulva may be valid.[24] In the far north, red fox fossils have been found inSangamonian Stage deposits near theFairbanks District,Alaska, andMedicine Hat,Alberta. Fossils dating from the Wisconsinan are present in 25 sites acrossArkansas,California,Colorado,Idaho,Missouri,New Mexico,Ohio,[25]Tennessee,Texas,Virginia, andWyoming. Although they ranged far south during the Wisconsinan, the onset of warm conditions shrank their range toward the north, and they have only recently reclaimed their former North American ranges because of human-induced environmental changes.[26]Genetic testing indicates that two distinct red foxrefugia exist in North America, which have been separated since the Wisconsinan. The northern (or boreal) refugium occurs in Alaska and western Canada, and consists of the larger subspeciesV. v. alascensis,V. v. abietorum,V. v. regalis, andV. v. rubricosa. The southern (or montane) refugium occurs in the subalpine parklands and alpine meadows of the west, from theRocky Mountains to theCascades and theSierra Nevada ranges, consisting of the smaller subspeciesV. v. cascadensis,V. v. macroura,V. v. necator, andV. v. patwin. The latterclade has been isolated from all other red fox populations since at least the last glacial maximum and may have developed unique ecological or physiological adaptations.[23]

Although European foxes (V. v. crucigera) were introduced to parts of the United States in the 1900s, recent genetic research suggests that there are no European fox mitochondrial haplotypes present in any North American populations.[27] Additionally, introduced eastern North American red foxes have colonised most of inland California, fromSouthern California to theSan Joaquin Valley,Monterey and north-coastalSan Francisco Bay Area (including urbanSan 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 theSacramento Valley red fox (V. v. patwin) only in a narrow hybrid zone.[28] In addition, no evidence is seen of interbreeding of eastern American red foxes in California with the montaneSierra Nevada red fox (V. v. necator) or other populations in the Intermountain West (between theRocky Mountains to the east and the Cascade and Sierra Nevada Mountains to the west).[29]

Subspecies

[edit]

The third edition ofMammal 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 throughmitochondrialhaplotype studies.[30] Castello (2018) recognized 30 subspecies of the Old World red fox and nine subspecies of theNorth American red fox as valid.[31]

It is known that substantialgene pool mixing occurs between different subspecies. For example, British red foxes havecrossbred extensively with red foxes imported from Germany, France, Belgium, Sardinia and possibly Siberia and Scandinavia.[32]: 140  However, genetic studies suggest very little differences between red foxes sampled across Europe.[33][34] 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.[35]

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 subspeciesV. v. griffithi,V. v. pusilla, andV. 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]

  • Skull of a northern fox
    Skull of a northern fox
  • Skull of a southern grey desert fox
    Skull of a southern grey desert fox
SubspeciesTrinomial authorityTrinomial authority (year)DescriptionRangeSynonyms
Scandinavian red fox
V. v. vulpes(nominate subspecies)

Linnaeus1758A large subspecies measuring 70–90 cm in length and weighing 5–10 kg; the maximum length of the skull for males is 163.2 mm. The fur is bright red with a strongly developed whitish and yellow ripple on the lower back.[7]Scandinavia and the northern and middle (forest) districts of the European part of theformer Soviet Union, southwards to the forest-steppe and eastwards approximately to theUrals and probably Central and Western Europe
  • alopex (Linnaeus, 1758)
  • communis (Burnett, 1829)
  • lineatus (Billberg, 1827)
  • nigro-argenteus (Nilsson, 1820)
  • nigrocaudatus (Billberg, 1827)
  • septentrionalis (Brass, 1911)
  • variegates (Billberg, 1827)
  • vulgaris (Oken, 1816)
British Columbia red fox[36]
V. v. abietorum

Merriam1900Generally similar toV. v. alascensis, but with a lighter, longer and more slender skull[37]SouthernYukon, theNorth-West Territories, northernAlberta, the interior ofBritish Columbia and in the adjoining coastal southeast Alaska (US).[38]sitkaensis (Brass, 1911)
Northern Alaskan fox
V. v. alascensis

Merriam1900A large, long-tailed, small-eared form with golden-fulvous fur[37]TheAndreafsky Wilderness, Alaska, U.S.
Eastern Transcaucasian fox
V. v. alpherakyi

Satunin1906A small subspecies weighing 4 kg; its maximum skull length is 132–39 mm in males and 121–26 mm in females. The fur is rusty grey or rusty brown, with a brighter rusty stripe along the spine. The coat is short, coarse and sparse.[7]Geok Tepe,Aralsk,Kazakhstan
Anatolian fox
V. v. anatolica

Thomas1920İzmir, theAegean Region,Turkey
Arabian red fox
V. v. arabica

Thomas1920Dhofar and theHajar Mountains,Oman
Atlas fox
V. v. atlantica

Wagner1841TheAtlas Mountains,Mila Province,Algeriaalgeriensis (Loche, 1858)
Labrador fox
V. v. bangsi

Merriam1900Similar toV. v. fulva, but with smaller ears and less pronounced black markings on the ears and legs.[37]L'Anse au Loup, theStrait of Belle Isle,Labrador,Canada
Barbary fox
V. v. barbara
Shaw1800TheBarbary Coast, northwestern Africaacaab (Cabrera, 1916)
Anadyr fox
V. v. beringiana

Middendorff1875A large subspecies; it is the most brightly coloured of the Old World red foxes, the fur being saturated bright-reddish and almost lacking the bright ripple along the back and flanks. The coat is fluffy and soft.[7]The shores of theBering Strait, northeasternSiberiaanadyrensis (J. A. Allen, 1903)

beringensis (Merriam, 1902)
kamtschadensis (Brass, 1911)
kamtschatica (Dybowski, 1922)
schantaricus (Yudin, 1986)

Cascade red fox
V. v. cascadensis

Merriam1900A short-tailed, small-toothed subspecies with yellow rather than fulvous fur; it is the subspecies most likely to produce "cross" colour morphs.[37]TheCascade Mountains,Skamania County,Washington, U.S.
North Caucasian fox
V. v. caucasica

Dinnik1914A large subspecies; its coat is variable in colour, ranging from reddish to red-grey and nearly grey. The fur is short and coarse. This subspecies could be a hybrid caused by mixing the populations ofV. v. stepensis andV. v. karagan.[7]NearVladikavkaz, theCaucasus,Russia
European fox
V. v. crucigera

Bechstein1789A medium-sized subspecies; its yellowish-fulvous or reddish-brown pelt lacks the whitish shading on the upper back. The tail is not grey, as in most other red fox subspecies.[39] It is primarily distinguished fromV. v. vulpes by its slightly smaller size, distinctly smaller teeth and widely spaced premolars. Red foxes present in Great Britain (and therefore Australia) are usually ascribed to this subspecies, though many populations there display a great degree of tooth compaction not present in continental European red fox populations.[9]All of Europe except Scandinavia, the Iberian Peninsula and some islands of the Mediterranean Sea; introduced to Australia and North Americaalba (Borkhausen, 1797)

cinera (Bechstein, 1801)
diluta (Ognev, 1924)
europaeus (Kerr, 1792)
hellenica (Douma-Petridou and Ondrias, 1980)
hypomelas (Wagner, 1841)
lutea (Bechstein, 1801)
melanogaster (Bonaparte, 1832)
meridionalis (Fitzinger, 1855)
nigra (Borkhausen, 1797)
stepensis (Brauner, 1914)

Trans-Baikal fox
V. v. daurica
Ognev1931A large subspecies; the colour along its spine is light, dull yellowish-reddish with a strongly developed white ripple and greyish longitudinal stripes on the anterior side of the limbs. The coat is coarse but fluffy.[7]Kharangoi, 45 km west ofTroizkosavsk, Siberiaussuriensis (Dybowski, 1922)
Newfoundland fox
V. v. deletrix
Bangs1898A very pale-coloured form; its light, straw-yellow fur deepens to golden yellow or buff-fulvous in some places. The tail lacks the usual black basal spot. The hind feet and claws are very large.[37]St. George's Bay,Newfoundland, Canada
Ussuri fox
V. v. dolichocrania
Ognev1926Sidemi, southernUssuri, southeastern Siberiaognevi (Yudin, 1986)
V. v. dorsalisJ. E. Gray1838
Turkmenian fox
V. v. flavescens

J. E. Gray1838A small subspecies with an infantile-looking skull and an overall grey-coloured coat; its body length is 49–57.5 cm and it weighs 2.2–3.2 kg.[7]NorthernIrancinerascens (Birula, 1913)
splendens (Thomas, 1902)
American red fox
V. v. fulva

Desmarest1820This is a smaller subspecies thanV. v. vulpes, with a smaller, sharper face, a shorter tail, a lighter pelt more profusely mixed with whitish and darker limbs.[37]Eastern Canada and the eastern U.S.pennsylvanicus (Rhoads, 1894)
Afghan red fox
V. v. griffithi

Blyth1854Slightly smaller thanV. v. montana; it has a more extensively hoary and silvered pelt.[40]: 121 Kandahar,Afghanistanflavescens (Hutton, 1845)
Kodiak fox
V. v. harrimani

Merriam1900This large subspecies has an enormous tail and coarse, wolf-like fur on the tail and lower back. The hairs on the neck and shoulders are greatly elongated and form a ruff.[37]Kodiak Island, Alaska, U.S.
South Chinese fox
V. v. hoole
R. Swinhoe1870[41]NearAmoy,Fukien, southern Chinaaurantioluteus (Matschie, 1907)
lineiventer (R. Swinhoe, 1871)
Sardinian fox
V. v. ichnusae
Miller1907A small subspecies with proportionately small ears.[39]Sarrabus,Sardinia,Italy; may have been introduced to theEnglish Midlands[32]: 6 
Cyprus fox
V. v. indutus
Miller1907Cyprus
Yakutsk fox
V. v. jakutensis

Ognev1923This subspecies is large, but smaller thanV. v. beringiana. The back, neck and shoulders are brownish-rusty, while the flanks are bright ocherous reddish-yellow.[7]Thetaiga south ofYakutsk, eastern Siberiasibiricus (Dybowski, 1922)
Japanese fox
V. v. japonica

Ognev1923Japan, except forHokkaido
Karaganka fox
V. v. karagan

Erxleben1777A smaller subspecies thanV. v. vulpes; its fur is short, coarse and of a light sandy-yellow or yellowish-grey colour.[7]TheKirghiz Steppes,Khirgizia, Russiaferganensis (Ognev, 1926)

melanotus (Pallas, 1811)
pamirensis (Ognev, 1926)
tarimensis (Matschie, 1907)

Kenai Peninsula fox
V. v. kenaiensis
Merriam1900One of the largest North American subspecies; it has softer fur thanV. v. harrimani.[37]TheKenai Peninsula, Alaska, U.S.
Transcaucasian montane fox
V. v. kurdistanica

Satunin1906A form intermediate in size betweenV. v. alpheryaki andV. v. caucasica; its fur is pale yellow or light grey, sometimes brownish-reddish and is fluffier and denser than that of the other two Caucasian red fox subspecies.[7]Northeastern Turkeyalticola (Ognev, 1926)
Wasatch Mountains fox
V. v. macroura

Baird1852This fox is similar toV. v. fulvus, but with a much longer tail, larger hind feet and more extensive blackening of the limbs.[37]Named for theWasatch Mountains near theGreat Salt Lake,Utah, found in theRocky Mountains fromColorado andUtah, westernWyoming andMontana throughIdaho north to southernAlberta
Hill fox
V. v. montana

Pearson1836This subspecies is distinguished fromV. v. vulpes by its smaller size, proportionately smaller skull and teeth and coarser fur. The hairs on the sole of the feet are copiously mixed with softer, woolly hairs.[40]: 111 TheHimalayas and northernIndian subcontinentalopex (Blanford, 1888)

himalaicus (Ogilby, 1837)
ladacensis (Matschie, 1907)
nepalensis (J. E. Gray, 1837)
waddelli (Bonhote, 1906)

Sierra Nevada red fox or High Sierra fox
V. v. necator

Merriam1900Externally similar toV. v. fulvus; it has a short tail, but cranially it is more likeV. v. macroura[37]TheHigh Sierra, California
Nile fox
V. v. niloticus

E. Geoffroy Saint-Hilaire1803A small subspecies; it measures 76.7–105.3 cm in body length, 30.2–40.1 cm in tail length and weighs 1.8–3.8 kg. It is ruddy to grey-brown above and darker on the back of the neck. The flanks are greyer and tinged with buff.[42] It is larger thanV. v. arabica andV. v. palaestina.[43]Egypt andSudanaegyptiacus (Sonnini, 1816)

anubis (Hemprich and Ehrenberg, 1833)
vulpecula (Hemprich and Ehrenberg, 1833)

Turkestan fox
V. v. ochroxantha
Ognev1926Aksai,Semirechye, eastern RussianTurkestan, Kirgizia
Palestinian fox
V. v. palaestina

Thomas1920Ramleh, nearJaffa,Israel
Korean fox
V. v. peculiosa

Kishida1924Northeastern China, Southeastern Russia, andKoreakiyomassai (Kishida andMori, 1929)
White-footed fox
V. v. pusilla

Blyth1854Slightly smaller thanV. v. griffithii;[40]: 123  it closely resembles theBengal fox (V. bengalensis) in size, but is distinguished by its longer tail and hind feet.[40]: 129 TheSalt Range,Punjab, Pakistanleucopus (Blyth, 1854)

persicus (Blanford, 1875)

Northern plains fox
V. v. regalis

Merriam1900The largest North American red fox subspecies; it has very large and broad ears and a very long tail. It is a golden-yellow colour with pure black feet.[37]TheElk River,Sherburne County,Minnesota, US
Nova Scotia fox
V. v. rubricosa
Bangs1898A large-sized subspecies with a large, broad tail and larger teeth and rostrum thanV. v. fulvus; it is the deepest-coloured subspecies.[37]Digby County,Nova Scotia, Canadabangsi (Merriam, 1900)

deletrix (Bangs, 1898)
rubricos (Churcher, 1960)
vafra (Bangs, 1897)

Ezo red fox
V. v. schrencki

Kishida1924Sakhalin, Russia andHokkaido, Japan
Iberian fox
V. v. silacea

Miller1907Though equal in size toV. v. vulpes, it has smaller teeth and more widely spaced premolars. The fur is dull buff without any yellowish or reddish tints. The hindquarters are frosted with white and the tail is clear grey in colour.[44]TheIberian Peninsula
Kurile Islands fox
V. v. splendidissima
Kishida1924The northern and centralKurile Islands, Russia
Steppe red fox
V. v. stepensis
Brauner1914This subspecies is slightly smaller and more lightly coloured thanV. v. crucigera, with shorter, coarser fur. Specimens from the Crimean Mountains have brighter, fluffier and denser fur.[7]The steppes nearKherson, Ukrainekrymeamontana (Brauner, 1914)

crymensis (Brauner, 1914)

Tobolsk fox
V. v. tobolica
Ognev1926This large subspecies has yellowish-rusty or dirty-reddish fur with a well-developed cross and often a black area on the belly. The coat is long and fluffy.[7]Obdorsk,Tobolsk, Russia
North Chinese fox
V. v. tschiliensis

Matschie1907Slightly larger thanV. v. hoole, but unlike other Chinese red foxes, it closely approachesV. v. vulpes in size.[45]Beijing andHebei province, Chinahuli (Sowerby, 1923)

Description

[edit]

Build

[edit]
Red fox (left) andcorsac fox (right) yawning

The red fox has an elongated body and relatively short limbs. The tail, which is longer than half the body length[7] (70 percent of head and body length),[46] is fluffy and reaches the ground when in a standing position. Their pupils are oval and vertically oriented.[7]Nictitating membranes are present, but move only when the eyes are closed. The forepaws have five digits, while the hind feet have only four and lackdewclaws.[9] They are very agile, being capable of jumping over 2 m (6 ft 7 in) high fences, and swim well.[47] Vixens normally have four pairs ofteats,[7] though vixens with seven, nine, or ten teats are not uncommon.[9] Thetestes of males are smaller than those of Arctic foxes.[7]

Theirskulls are fairly narrow and elongated, with smallbraincases. Theircanine 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 hardpalates, as well as having larger canines.[7] Their skulls are distinguished from those of dogs by their narrowermuzzles, less crowdedpremolars, more slender canine teeth, and concave rather than convex profiles.[9]

Dimensions

[edit]

Red foxes are the largest species of the genusVulpes.[48] However, relative to their size, red foxes are much lighter than similarly sized dogs in the genusCanis. For example, their limb bones weigh 30 percent less per unit area of bone than expected for similarly sized dogs.[49]: 122  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 tods.[50][51] 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.[49]: 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.[52] 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 inAberdeenshire, Scotland, in early 2012.[53]

Fur

[edit]
A red fox in its winter coat in the Rocky Mountains of Colorado, U.S.

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.[54] Among northern foxes, the North American varieties generally have the silkiestguard hairs,[13]: 231  while most Eurasian red foxes have coarser fur.[13]: 235  The fur on areas such as the head and lower legs, known as "thermal windows", is kept short and dense all year round, while the fur on other areas changes with the seasons. Foxes actively control the peripheralvasodilation and peripheralvasoconstriction in these areas to regulate heat loss.[55] There are three maincolour morphs; red, silver/black and cross (seeMutations).[46] In the typical red morph, their coats are usually bright reddish-rust colour with yellowish tints. Along the spine, there is a stripe of weak, diffuse patterns of brown, reddish and chestnut hairs. Two additional stripes run down the shoulder blades and, together with the spinal stripe, form a cross. The lower back is often a mottled silvery colour. The flanks are lighter coloured than the back, while the chin, lower lips, throat and front of the chest are white. The remaining lower surface of the body is dark, brown or reddish.[54] During lactation, the belly fur of vixens may turn brick red.[9] The upper parts of the limbs are rusty red, while the paws are black. The front of the face and the upper neck are bright brownish-red, while the upper lips are white. The backs of the ears are black or brownish-reddish, while the inner surface is whitish. The top of the tail is brownish-reddish, but lighter in colour than the back and flanks. The underside of the tail is pale grey with a straw-coloured tint. A black spot, the location of thesupracaudal gland, is usually present at the base of the tail. The tip of the tail is white.[56]

Colour morphs

[edit]
Various red fox colour morphs

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

Colour morphImageDescription
RedThe typical colouration (seefur above)
SmokeyThe 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]
CrossThe 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-brownThe 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]
SilverThe 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]
PlatinumDistinguished from the silver colour morph by its pale, almost silvery-white fur with a bluish cast[13]: 251 
Amber
SamsonDistinguished by its woolly pelt, which lacks guard hairs[13]: 230 

Senses

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Red foxes havebinocular vision,[9] but their sight reacts mainly to movement. They have acuteauditory perception, being able to hearblack grouse changing roosts at 600 paces,crows flying at 0.25–0.5 km (0.16–0.31 mi) and mice squeaking at about 100 m (330 ft).[57] They can locate sounds with an accuracy of one degree at frequencies of 700–3,000 Hz, though less accurately at higher frequencies.[47] Their sense of smell is good, but weaker than that of dogs specialised in scent work.[57]

Scent glands

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Red foxes have a pair ofanal sacs lined with sebaceous glands that open through a single duct.[58] The size and volume of the anal sacs increases with age, ranging in size from 5–40mm in length, 1–3mm in diameter, and with a capacity of 1–5 mL.[59] The anal sacs act as fermentation chambers in whichaerobic andanaerobic bacteria convertsebum into odorous compounds, includingaliphatic acids. The oval-shapedcaudal gland is 25 mm (0.98 in) long and 13 mm (0.51 in) wide, and reportedly smells ofviolets.[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

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V. v. crucigera, Slovakia
Multi-coloured North American red fox

The red fox is a widespread species. Its range covers nearly 70,000,000 km2 (27,000,000 sq mi), extending as far north as theArctic Circle. It is found throughout 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 from the Arctic islands,[disputeddiscuss] the most northerly parts of centralSiberia, and in extreme deserts.[1]It is not present inNew Zealand and is classed as a "prohibited new organism" under theHazardous Substances and New Organisms Act of 1996, which does not allow its import.[60]

Australia

[edit]
Main article:Feral foxes in Australia

In Australia, estimates in 2012 indicated that there were more than 7.2 million red foxes,[61] with a range extending throughout most of the continental mainland.[49]: 14  They were introduced Australia in the 1830s and 1840s by settlers in the British colonies ofVan Diemen's Land (as early as 1833) and thePort Phillip District of New South Wales (as early as 1845). The settlers wanted to foster the traditional English sport offox hunting. However, a permanent red fox population did not become established on the island ofTasmania, and it is widely believed that foxes were outcompeted by theTasmanian devil.[62] On the mainland, however, the species was successful as anapex predator. The fox is generally less common in areas where thedingo is more prevalent, but it has achievedniche differentiation with both theferal dog and theferal cat, primarily through its burrowing behaviour. 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 familyPotoroidae, including thedesert rat-kangaroo.[63] The spread of red foxes across the southern part of the continent has coincided with the spread ofrabbits in Australia, and corresponds with declines in the distribution of several medium-sized ground-dwelling mammals, includingbrush-tailed bettongs,burrowing bettongs,rufous bettongs,bilbies,numbats,bridled nail-tail wallabies andquokkas.[64] Most of those species are now limited to areas (such as islands) where red foxes are absent or rare. Local fox eradication programs exist, although elimination has proven difficult due to the fox's denning behaviour and nocturnal hunting, so the focus is on management, including the introduction of state bounties.[65] According to the Tasmanian government, red foxes were accidentally introduced to the previously fox-free island of Tasmania in 1999 or 2000, posing a significant threat to native wildlife, including theeastern bettong, and an eradication program was initiated, conducted by the TasmanianDepartment of Primary Industries and Water.[66]

Sardinia, Italy

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Current range of the red fox in Italy (orange) alongside the range of theEuropean jackal andItalian wolf

The origin of theichnusae subspecies inSardinia, Italy is uncertain, as it is absent from Pleistocene deposits in their current homeland. It is possible it originated during theNeolithic 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.[22]

Behaviour and ecology

[edit]

Social and territorial behaviour

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Red fox pressed against the trunk of a pine tree inIlmatsalu, Estonia
Red foxesgrooming each other
A pair of European red foxes (V. v. crucigera) at theBritish Wildlife Centre, Surrey, England
A red foxmarking its territory

Red foxes either establish stable home ranges within particular areas or are itinerant with no fixed abode.[49]: 117  They use theirurine to mark their territories.[67][68] A male fox raises one of his hind legs, spraying his urine forward in front of him. In contrast, a female fox squats down, spraying her urine onto the ground between her hind legs.[69] Urine is also used to mark emptycache sites where found food is stored. This serves as a reminder not to waste time investigating them.[49]: 125 [70][71] Males generally have higher urine marking rates during late summer and autumn, but the rest of the year the rates between male and female are similar.[72] The use of up to 12 different urination postures allows them to precisely control the position of the scent mark.[73]

Red foxes live in family groups that share a territory. In favourable habitats and/or areas with low hunting pressure, a range of subordinate foxes may be present. There may be one or two subordinate foxes, or sometimes up to eight, in one territory. These subordinates may formerly have beendominant animals, but they are mostly young from the previous year and act as helpers in rearing the breeding vixen's kits. Alternatively, their presence has been explained as a response to temporary food surpluses unrelated to assisting reproductive success. Non-breeding vixens guard, play with, groom, provision and retrieve kits,[9] an example ofkin selection. Red foxes may leave their families once they reach adulthood if they have a high chance of winning of their own. Otherwise, they will stay with their parents, postponing their own reproduction in the process.[49]: 140–141 

Reproduction and development

[edit]
Further information:Mating behavior of melanistic red foxes
Red fox kits
Red foxesmating

Red foxes reproduce once a year in spring. Two months prior tooestrus (typically December), the reproductive organs of vixens change shape and size. By the time they enter their oestrus period, theiruterine horns double in size, and theirovaries grow 1.5–2 times larger.Sperm formation in males begins in August–September, with the testicles attaining their greatest weight in December–February.[74] The vixen's oestrus period lasts three weeks,[9] during which the dog-foxes mate with the vixens for several days, often in burrows. The male'sbulbus glandis enlarges duringcopulation,[10] forming acopulatory tie which may last for more than an hour.[9] Thegestation period lasts 49–58 days.[74] Though foxes are largelymonogamous,[75] DNA evidence from one population indicated large levels ofpolygyny,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 been reported.[76] Large litters are typical in areas where fox mortality is high.[49]: 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.[77] Mothers remain with the kits for 2–3 weeks as they are unable tothermoregulate. 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.[32]: 21–22  If the mother dies before the kits are independent, the father takes over as their provider.[32]: 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.[77] Their eyes are blue at birth, but change to amber at 4–5 weeks. Their coat begins to change colour at three weeks of age, when a black streak appears around the eyes. By one month of age, red and white patches appear on their faces. During this time, their ears become erect and their muzzles elongate.[9] Kits begin to leave their dens and experiment with solid food provided by their parents at the age of 3–4 weeks. Thelactation period lasts 6–7 weeks.[77] 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.[77] Some vixens may reachsexual maturity at the age of 9–10 months, thus bearing their first litters at one year of age.[74] In captivity, theirlongevity can be as long as 15 years, though in the wild they typically do not survive past 5 years of age.[78]

Denning behaviour

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Side and above view of a red fox den

Outside of thebreeding season, most red foxes prefer to live in the open, in densely vegetated areas, although they may enter burrows to escape bad weather.[9] Their burrows are often dug on hill or mountain slopes, in ravines and on steep riverbanks, as well as in ditches, depressions and gutters, and in rock clefts and neglected human environments. Red foxes prefer to dig their burrows in well-drained soil. Dens built among tree roots can last for decades, whereas those dug in steppe regions only last several years.[7] They may permanently abandon their dens duringmange outbreaks, possibly as a defence mechanism against the spread of disease.[9] In the Eurasian desert regions, foxes may use theburrows of wolves, porcupines and other large mammals, as well as those dug by gerbil colonies. Compared to burrows constructed by Arctic foxes, badgers, marmots and corsac foxes, red fox dens are not overly complex. Red fox burrows consist of a den and temporary burrows. The latter comprise only a small passage or cave for concealment. The main entrance of the burrow leads downwards (40–45°) and opens into a den, from which numerous side tunnels branch off. The depth of the burrow ranges from 0.5–2.5 m (1 ft 8 in – 8 ft 2 in), rarely extending toground water. The main passage can reach 17 m (56 ft) in length, standing an average of 5–7 m (16–23 ft). In spring, red foxes clear their dens of excess soil by making rapid movements with their forepaws and hind legs. They throw the discarded soil up to 2 m (6 ft 7 in) from the burrow. Once kits are born, the discarded debris is trampled to form an area where the kits can play and receive food.[7] They may share their dens with woodchucks[10] or badgers.[7] Unlike badgers, which fastidiously clean their earths and defecate inlatrines, red foxes habitually leave pieces of prey around their dens.[32]: 15–17  The average sleep time of a captive red fox is 9.8 hours per day.[79]

Communication

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

The body language of red foxes consists of movements of the ears and tail, as well as postures. Their body markings emphasise certain gestures. These postures can be categorised as either aggressive/dominant or fearful/submissive. Some postures may blend the two together.[49]: 42–43 Inquisitive foxes will rotate and flick their ears while they sniff. Playful individuals will perk their ears up and stand on their hind legs. Male foxes will turn their ears outward and raise their tails horizontally with the tips raised upwards when courting females or after successfully evicting intruders. 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, they adopt a similar posture, but without arching their backs or curving their bodies. Submissive foxes approach dominant animals with their muzzles reaching up in greeting, in a low posture. When two evenly matched foxes confront each other over food, they approach side by side and push against each other's flanks. They betray a mixture of fear and aggression through their actions: they lash their tails and arch their backs, but do not crouch or pull their ears back and flatten them against their skulls. When launching an assertive attack, red foxes approach directly, with their tails held high and their ears rotated sideways.[49]: 43  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]

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

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Red foxes have a wide vocal range, and produce different sounds spanning fiveoctaves which blend into each other.[49]: 28  Recent analyses have identified 12 different sounds produced by adults and 8 by kits.[9] The majority of sounds can be categorised as either "contact" or "interaction" calls. The former vary according to the distance between individuals, while the latter vary according to the level of aggression.[49]: 28 

  • Contact calls: The most commonly heard contact call is a barking sound consisting of three to five syllables, transcribed as "wow wow wow", 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 oftawny owls). The "wow wow wow" call varies from one fox to another. 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.[49]: 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 aratchet, called "gekkering". Gekkering occurs mostly during the courting season from rival males or vixens rejecting advances.[49]: 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.[49]: 28 

Diet and hunting

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A red fox with acoypu

The red fox is anomnivore with a highly varied diet.[80][81]It consumes over 300 animal species including small rodents likemice,ground squirrels,hamsters,gerbils,woodchucks,muskrats,voles,pocket gophers,deer mice,leporids,porcupines,raccoons,opossums, birds withpasseriformes,galliformes andwaterfowl predominating,reptiles,insects andinvertebrates.[7][10] It attacks young or smallungulates on rare occasions.[7] It typically targets mammals with a weight of up to 3.5 kg (7.7 lb) and requires 500 g (18 oz) of food daily.[47]It hunts its preferred prey when it is abundant, but it can still survive on different prey, as well as seeds and fruit, when it is not available.[82] It occasionally commits acts ofsurplus killing; during one breeding season, four red foxes were recorded to have killed around 200black-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][49]: 164  In some areas, fruit amounts to 100% of its diet in autumn, includingblueberry,blackberry,raspberry,cherry,persimmon,mulberry,apple,plum,grape andacorn. Other plant material includesgrass,sedge andtuber.[10]

In southern Italy, red foxes were observed eatingpond slider eggs excavated from nests.[83] Camera traps at an Australianrookery documented red foxes preying onflatback sea turtle eggs.[84]

Visual cues are the most important ones for the hunting behaviour of the red fox.[85]It prefers to hunt in the early morning hours before sunrise and late evening.[7] Successful hunting in long vegetation or under snow appears to involve its alignment with the Earth'smagnetic field.[86]It typically forages alone, but occasionally aggregates in resource-rich environments.[78] When hunting mouse-like prey, it first pinpoints the prey's location by sound, then leap, sailing high above the quarry, steering in mid-air with its tail before landing on target up to 5 m (16 ft) away.[1] It feeds oncarrion in the late evening hours and at night.[7] It seem to dislike thetaste ofmoles, but nonetheless catches them alive and presents them to its young for playing.[49]: 41  It is possessive of its food and defends its catches even from dominant animals.[49]: 58 

Enemies and competitors

[edit]
A red fox confronting a gray fox

The red fox typically dominates other fox species. Where the red fox andArctic fox aresympatric, the Arctic fox feeds onlemmings and flotsam rather than voles, as favoured by red foxes. Both species kill each other's kits, given the opportunity.[7] The southern range of the Arctic fox is limited by the presence of the red fox; both were introduced to almost every island from theAleutian Islands to theAlexander Archipelago during the 1830s–1930s by fur companies. The red fox invariably displaced the Arctic fox, with one male red fox having been reported to have killed off all resident Arctic foxes on a small island in 1866.[49]: 85 

The red fox is a competitor of thecorsac fox, as they hunt the same prey all year. The red fox is stronger, 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. The corsac fox seems to only outcompete the red fox in semi-desert and steppe areas.[7][87] In Israel, the Blanford's fox escapes competition with the red fox by being restricted to rocky cliffs and actively avoiding the open plains inhabited by the red fox.[49]: 84–85  The red fox dominatekit andswift 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] Thegray fox dominates the red fox wherever their ranges meet. Historically, interactions between the two species were rare, as the gray fox favoured heavily wooded or semiarid habitats as opposed to the open and mesic ones preferred by the red fox. However, interactions have become more frequent due to deforestation, allowing the red fox to colonise gray fox-inhabited areas.[10]

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

Thewolf is known to kill and eat the red fox in disputes over carcasses.[7][88] In North America where the red fox andcoyote are sympatric, theirhome ranges do not overlap, and 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.[89] In Israel, the red fox shares habitat with thegolden jackal, and where their ranges meet, they compete due to near-identical diets. The red fox ignores golden jackal scents or tracks and avoids close physical proximity; in areas where the golden jackal is abundant, the red fox population decreases significantly, apparently because ofcompetitive exclusion.[90] However, multiple red foxes interacting peacefully with a golden jackal have been observed in southwestern Germany.[91]

Agolden eagle feeding on a red fox

The red fox is dominant over theraccoon dog and will sometimes kill its kits or bite adults to death. There are documented cases of red foxes killing raccoon dogs by entering their dens. Both species compete for prey similar in size to mice. This competition peaks in early spring when food is scarce. InTatarstan, 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]

The red fox occasionally kills smallmustelids likeleast weasel,[10]stone marten,[92]pine marten,stoat,Siberian weasel,European polecat and youngsables.Eurasian badgers may live alongside red foxes in isolated sections of large burrows.[7] The two species may tolerate each other out ofmutualism: red foxes provide Eurasian badgers with food scraps, while Eurasian badgers keep the shared burrow clean.[32]: 15  However, there are known cases of Eurasian badgers driving vixens from their dens and destroying their litters without eating them.[7]Wolverines may kill red foxes, often while the latter is sleeping or near carrion.[7]: 546  In turn, red foxes may kill young wolverines.[7]

The red fox competes with thestriped hyena over large carcasses. The striped hyena's stronger jaws can easily tear open flesh that is too tough for the red fox. When harassing striped hyenas, it avoids attacks by outrunning them. Sometimes, red foxes seem to deliberately torment striped hyenas even when food is not at stake. However, some red foxes misjudge their attacks and are killed.[49]: 77–79  The remains of red foxes are often found in striped hyena dens, and striped hyenas steal red foxes from traps.[7]

The red fox may be preyed upon by theleopard,caracal andEurasian lynx. The Eurasian lynx chases the red fox into deep snow, where its long legs and larger paws gives it an advantage over the red fox, especially when the depth of the snow exceeds one meter.[7] In theVelikoluksky District in Russia, the red fox is absent or seen only occasionally where the Eurasian lynx established permanent territories.[7] In North America, the red fox is predated by thecougar,Canada lynx andbobcat.[46]

The red fox competes withcommon buzzard (Buteo buteo) andnorthern goshawk (Accipiter gentilis) and even steals their kills.[93] Thegolden eagle (Aquila chrysaetos) regularly takes young red foxes and preys on adults if needed.[94] Other large eagles such as thewedge-tailed eagle (Aquila audax),eastern imperial eagle (Aquila heliaca),white-tailed eagle (Haliaeetus albicilla) andsteller's sea eagle (Haliaeetus pelagicus) occasionally kill red foxes.[95][96][97] Large owls such as theEurasian eagle-owl (Bubo bubo) andsnowy owl (Bubo scandiacus) prey on young foxes and adults on occasions.[98][99]

Diseases and parasites

[edit]
A European fox (V. v. crucigera) withmange

Red foxes are the most importantrabies vector in Europe. InLondon,arthritis is common in foxes, being particularly frequent in the spine.[9] Foxes may be infected withleptospirosis andtularemia, though they are not overly susceptible to the latter. They may also fall ill fromlisteriosis andspirochetosis, as well as acting asvectors in spreadingerysipelas,brucellosis and tick-borneencephalitis. A mysterious fatal disease nearLake Sartlan in theNovosibirsk Oblast occurred among red foxes, but the cause was unclear. Individual cases of foxes infected withYersinia pestis are known.[7]

Red foxes are not readily prone to infestation withfleas. Species likeSpilopsyllus cuniculi are probably only caught from the fox's prey species, while others likeArchaeopsylla erinacei are caught whilst traveling. Fleas that feed on red foxes includePulex irritans,Ctenocephalides canis andParaceras melis.Ticks such asIxodes ricinus andI. hexagonus are not uncommon in red foxes, and are typically found on nursing vixens and kits still in their earths. ThelouseTrichodectes vulpis specifically targets red foxes, but is found infrequently. ThemiteSarcoptes scabiei is the most important cause ofmange in red foxes. It causes extensive hair loss, starting from the base of the tail and hindfeet, then the rump before moving on to the rest of the body. In the final stages of the condition, red foxes can lose most of their fur, 50% of their body weight and may gnaw at infected extremities. In theepizootic phase of the disease, it usually takes red foxes four months to die after infection. Other endoparasites includeDemodex folliculorum,Notoderes,Otodectes cynotis (which is frequently found in theear canal),Linguatula serrata (which infects the nasal passages) andringworms.[7]Up to 60helminth species are known to infect captive-bred foxes infur farms, while 20 are known in the wild. Severalcoccidian species of thegeneraIsospora andEimeria are also known to infect them.[7] The most commonnematode species found in red fox guts areToxocara canis andUncinaria stenocephala,Capillaria aerophila[100] andCrenosoma vulpis; the latter two infect their lungs and trachea.[101]Capillaria plica infects the red fox's bladder.Trichinella spiralis rarely affects them. The most commontapeworm species in red foxes areTaenia spiralis andT. pisiformis. Others includeEchinococcus granulosus andE. multilocularis. Eleventrematode species infect red foxes,[9] includingMetorchis conjunctus.[102] A red fox from was found to be a host of intestinal parasiticacanthocephalan worms,Pachysentis canicola inBushehr Province, Iran,[103]Pachysentis procumbens andPachysentis ehrenbergi in both in Egypt.[104]

Relationships with humans

[edit]

In folklore, religion and mythology

[edit]
Further information:Foxes in culture
Reynard the Fox in an 1869 children's book

The red fox features prominently in the folklore and mythology. InGreek mythology, theTeumessian fox,[105] or Cadmean vixen, was a gigantic fox that was destined never to be caught. The fox was one of the children ofEchidna.[106]

InCeltic mythology, the red fox is a symbolic animal. In theCotswolds,witches were thought to transform into foxes in order to stealbutter from their neighbours.[107] In later European folklore, the figure ofReynard the Fox symbolises trickery and deceit. He first appeared as a secondary character under the name of "Reinardus" in the 1150 poem "Ysengrimus", and then in Pierre Saint Cloud'sLe Roman de Renart in 1175. He made his debut in England inGeoffrey Chaucer'sThe Nun's Priest's Tale. Many of Reynard's adventures may stem from actual observations of fox behaviour: he is an enemy of the wolf and has a fondness for blackberries and grapes.[49]: 32–33 

Chinese folk tales tell of fox-spirits calledhuli jing that may have up to nine tails, orkumiho as they are known in Korea.[108] InJapanese mythology, thekitsune are fox-like spirits possessing magical abilities that increase with their age and wisdom. Foremost among these is the ability todisguise as human. While some folktales speak of kitsune employing this ability to trick others, other stories portray them as faithful guardians, friends, lovers and wives.[109] InArab folklore, the fox is considered a cowardly, weak, deceitful and cunning animal that feigns death by filling its abdomen with air to appear bloated, then lies on its side, awaiting the approach of unwitting prey.[43] The animal's cunning was noted by the authors of theBible who applied the word "fox" to false prophets in theBook of Ezekiel 13:4 and the hypocrisy ofHerod Antipas inLuke 13:32.[110]

The cunning Fox is a common character innative American mythology, where it is often portrayed as a constant companion toCoyote. Fox, however, is a deceitful companion who often steals Coyote's food. In theAchomawicreation myth, Fox and Coyote are the co-creators of the world, that leave just before the arrival of humans. TheYurok tribe believed that in a fit of anger, Fox captured theSun and tied him to a hill, causing him to burn a great hole in the ground. In anInuit story, Fox is portrayed as a beautiful woman who tricks a hunter into marrying her, only to resume her true form and leave after he offends her. AMenominee story tells of how Fox is an untrustworthy friend to Wolf.[111]

Hunting

[edit]
Further information:Fox hunting
Beagle and Fox (1885) byBruno Liljefors

The earliest historical records of fox hunting date back to the 4th century BC;Alexander the Great is known to have hunted foxes, and aseal dated from 350 BC depicts aPersian horseman in the process of spearing a fox.Xenophon, who believed that hunting was part of a cultured man's education, promoted killing foxes as pests because they distracted hounds from hares. TheRomans were hunting foxes by AD 80. During theDark Ages in Europe, foxes were considered a secondary quarry, but gradually grew in importance.Cnut the Great reclassified foxes as Beasts of the Chase, which was a lower category of quarry than Beasts of Venery. Foxes were gradually hunted less as vermin and more as Beasts of the Chase, to the point that by the late 1200s,Edward I had a royalpack of foxhounds and a specialised fox huntsman. In this period, foxes were increasingly hunted above ground with hounds, rather than underground with terriers.Edward, Second Duke of York assisted the climb of foxes as more prestigious quarries in hisThe Master of Game. By theRenaissance, fox hunting became a traditional sport of the nobility. After theEnglish 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 theCharlton Hunt Club in 1737). The popularity of fox hunting in Great Britain reached a peak during the 1700s.[49]: 21  Although already native to North America, red foxes from England were imported for sporting purposes to Virginia andMaryland in 1730 by prosperous tobacco planters.[112] These American fox hunters considered the red fox more sporting than the grey fox.[112]

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 eightmiles. '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.[112]

Red foxes are still widely persecuted as pests, with human-caused deaths being one of the main causes of mortality for 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);[113]New Mexico (US) 69 (1999–2000).[92]

Fur use

[edit]
Red fox pelts
A red fox in a fur farm inVörå, Finland

Red foxes are among the most importantfur-bearing animals harvested by thefur trade. Their pelts are used fortrimmings,scarfs,muffs,jackets andcoats. They are primarily used as trimming for both cloth coats and fur garments, including evening wraps.[13]: 229–230  The pelts of silver foxes are popular ascapes,[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.[32]: 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.[32]: 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.[92] Pelt prices are increasing, with 2012 North American wholesale auction prices averaging $39 and 2013 prices averaging $65.78.[114]

Red foxes in North America, particularly those in northern Alaska, are highly valued for their fur. They have guard hairs with a silky texture that allows unrestricted mobility after dressing. However, red foxes living in southern Alaska's coastal areas and the Aleutian Islands are an exception, as their pelts are extremely coarse and rarely fetch more than a third of the price of those from northern Alaska.[13]: 231  Compared to North American varieties, most European peltries have coarse-textured fur. The only exceptions are Nordic and Far Eastern Russian pelts, which are still inferior to North American pelts in terms of silkiness.[13]: 235 

Livestock and pet predation

[edit]
A carcass of a lamb near a red fox den
A red fox in aBirmingham garden investigating a rabbit hutch

Red foxes may occasionally prey on lambs. Usually but not always, the lambs targeted are physically weakened specimens. Lambs belonging to smaller breeds, such as theScottish Blackface, are more vulnerable than larger breeds, such as theMerino. Twins may be more vulnerable to red foxes than single lambs, as ewes cannot effectively defend both simultaneously. Crossbreeding small, upland ewes with larger, lowland rams can result in difficult and prolonged labour for ewes due to the heaviness of the resulting offspring, making them more vulnerable to predation by red foxes. Lambs born to first-time mothers (gimmers) are more often killed by red foxes than lambs born to experienced mothers, who stick closer to their young.[49]: 166–167 

Red foxes may prey ondomestic rabbits andguinea pigs if they are kept in open runs or are allowed to roam freely in gardens. This problem can be avoided by housing them in robust hutches and runs. Urban red foxes often encountercats and may feed alongside them. In physical confrontations, the cats usually have the upper hand. Cases where red foxes have killed cats usually involve kittens. While most red foxes do not prey on cats, some may do so and may treat them more as competitors rather than food.[49]: 180–181 

Taming and domestication

[edit]
Further information:Domesticated silver fox
A young boy holding a tame red fox kit

In their unmodified wild state, red foxes are generally unsuitable as pets.[115] Many supposedly abandoned kits are adopted by well-meaning people in spring, although it is unlikely that vixens would abandon their young. Actual orphans are rare, and those that are adopted are probably just kits that have strayed from their dens.[116] 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.[49]: 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.[49]: 61  They maintain their wild counterparts' strong instinct of concealment and may pose a threat to domestic birds, even when well-fed.[32]: 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.[32]: 132–133 

White to black individual red foxes have been selected and raised on fur farms as "silver foxes". In the second half of the 20th century, a lineage ofdomesticated silver foxes was developed byRussian geneticistDmitry 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 includedpiebald coats, floppy ears in kits and curled tails, similar to the traits that distinguish domestic dogs from grey wolves.[117]

Urban red foxes

[edit]

Distribution

[edit]

Red foxes have been extremely successful in colonising built-up environments, particularly lower-density suburbs,[47] although they have also been sighted in densely populated urban areas far from the countryside. Throughout the 20th century, they have established themselves in many Australian, European, Japanese and North American cities. The species first colonised British cities during the 1930s, enteringBristol and London during the 1940s, and later established themselves inCambridge andNorwich. In Ireland, they are now common in suburbanDublin. In Australia, red foxes were recorded inMelbourne as early as the 1930s, while inZurich, Switzerland, they only started appearing in the 1980s.[118]

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.[47] 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).[119]

In the UK, it is estimated that the number of urban foxes rose from 33,000 in 1995 to 150,000 foxes in 2017.[120] City-dwelling red foxes may scavenge food from litter bins and bin bags: research in 2025 suggests that human-generated food comprises 35% of urban fox diet, compared to just 6% for their rural counterparts.[121] As a result of this difference in diet between urban and rural populations, city-dwelling red foxes tend to grow larger than their rural counterparts.[122] It has been observed that urban foxes tend to have shorter and wider muzzles, smaller braincases and reduced sexual dimorphism relative to rural individuals: this is thought to be as a result of differing biomechanical demands of feeding or cognition between habitats.[123]

Behaviour

[edit]

Urban red foxes are most active at dusk and dawn, when they do most of their hunting and scavenging. They are rarely seen during the day, but can sometimes be spotted sunbathing on the roofs of houses or sheds. They will often make their homes in hidden, undisturbed spots in urban areas, as well as on the outskirts of cities, visiting at night to find food. They sleep in dens at night.

While urban red foxes can scavenge successfully in cities, eating anything that humans eat, some people deliberately leave food out for them, finding them endearing. Doing this regularly can attract red foxes to one's home, where they can become accustomed to human presence. They may even allow themselves to be approached or played with, particularly the young kits.[119]

Urban red fox control

[edit]

Urban red foxes can cause problems for local residents. They are known to steal chickens, rummage through rubbish bins and damage gardens. Most complaints about urban red foxes made to local authorities occur during the breeding season, which is from late January/early February to late April/early August.[119]

In the UK, hunting red foxes in urban areas is banned, and shooting them in an urban environment is not a suitable alternative. Trapping them appears to be a more viable alternative to hunting urban red foxes.[124] However, killing red foxes has little effect on their urban population; the foxes that are killed are quickly replaced by new kits during the breeding season or by other foxes moving into the territory of those that were killed. A more effective method of controlling red foxes in urban areas is to deter them from specific areas. Deterrents such as creosote, diesel oil or ammonia can be used for this purpose. Cleaning up and blocking access to dens can also discourage red foxes from returning to urban areas.[119]

Relationship between urban and rural red foxes

[edit]

In January 2014 it was reported that "Fleet", a relatively tame urban red fox tracked as part of a wider study by theUniversity of Brighton in partnership with the BBC TV seriesWinterwatch, had unexpectedly traveled 195 miles in 21 days from his neighbourhood inHove at the western edge ofEast Sussex across rural countryside as far as the town ofRye, near the eastern edge of the county. He was still continuing his journey when theGPS 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.[125][126]

  • An urban red fox crossing a city street in Denver, Colorado
    An urban red fox crossing a city street in Denver, Colorado
  • An urban red fox in a built-up area in London, England
  • 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 seriesWinterwatch

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  96. ^Utekhina, I.; Potapov, E. & McGrady, M.J. (2000). "Diet of the Steller's Sea Eagle in the northern Sea of Okhotsk". In Ueta, M. & McGrady, M.J. (eds.).First Symposium on Steller's and White-tailed Sea Eagles in East Asia. Tokyo, Japan: Wild Bird Society of Japan. pp. 71–92.
  97. ^Vrezec, A.; Bordjan, D.; Perušek, M. & Hudoklin, A. (2009)."Population and ecology of the White-tailed Eagle (Haliaeetus albicilla) and its conservation status in Slovenia".Denisia.27:103–114.
  98. ^Dixon, C. C. (1970)."Red Fox predated by Snowy Owl".Blue Jay.33 (2).doi:10.29173/bluejay3979.
  99. ^"Eurasian Eagle Owl (Bubo bubo)".The Owl Pages. 2015.Archived from the original on 20 November 2015. Retrieved10 September 2016.
  100. ^Lalošević, V.; Lalošević, D.; Čapo, I.; Simin, V.; Galfi, A. & Traversa, D. (2013)."High infection rate of zoonoticEucoleus aerophilus infection in foxes from Serbia".Parasite.20 (3): 3.doi:10.1051/parasite/2012003.PMC 3718516.PMID 23340229.
  101. ^Mechouk, N.; Deak, G.; Ionică, A. M.; Toma, C. G.; Bouslama, Z.; Daniel Mihalca, A. (2023)."First report ofCrenosoma vulpis in Africa and Eucoleus aerophilus in Algeria".International Journal for Parasitology: Parasites and Wildlife.20:187–191.doi:10.1016/j.ijppaw.2023.03.003.PMC 10023903.PMID 36941973.
  102. ^Smith, H. J. (1978)."Parasites of red foxes in New Brunswick and Nova Scotia".Journal of Wildlife Diseases.14 (3):366–370.doi:10.7589/0090-3558-14.3.366.PMID 691132.
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  105. ^Ancient Greek:Τευμησ(σ)ία ἀλώπηξ (Teumēs(s)íā alôpēx),gen.: Τευμησίας ἀλώπεκος, also known as ἀλώπηξ τῆς Τευμησσοῦ "fox ofTeumessos"; Teumessos was an ancient city inBoeotia.
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Further reading

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External links

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Wikimedia Commons has media related tothe red fox.
Wikispecies has information related toRed fox.
Look upFox in Wiktionary, the free dictionary.
ExtantCarnivora species
Prionodon(Asiatic linsangs)
Pantherinae
Neofelis
Panthera
Felinaesensu stricto
Bay cat
lineage
Pardofelis
Catopuma
Caracal
lineage
Caracal
Leopardus
Lynx
Puma
lineage
Acinonyx
Puma
Leopard cat
lineage
Prionailurus
Felis
Viverroidea
    • see below↓
Hemigalinae
Paradoxurinae
Paradoxurus
Viverrinaesensu lato
Viverrinae
sensu stricto
Viverra
Poiana
(African linsangs)
Genetta
(genets)
Herpestoidea
    • see below↓
Hyaenidae
(hyenas)
Proteles
Hyaeninae
(bone-crushing hyenas)
Crocuta
Herpestidaesensu lato
Eupleridae
(Malagasy
carnivorans)
Euplerinae
(Malagasy civets)
Eupleres(falanoucs)
Galidiinae
(vontsira)
Galidictis
Salanoia
Suricata
Mungos
Helogale
Crossarchus
(kusimanses)
Urva
(Asian mongooses)
Bdeogale
Herpestes
(slender mongooses)
Urocyon
Nyctereutes
(raccoon dogs)
Vulpes
(truefoxes)
Speothos
Lycalopex
(South American foxes)
Lupulella
Lycaon
Canis
Ailuropoda
Tremarctos
Ursinae
Ursus
Mustelida
Pinnipedia(seals)
    • see below↓
Musteloidea
    • see below↓
Odobenidae
Callorhinus
(northernfur seals)
Otariinae
(sea lions)
Zalophus
Neophoca
Arctocephalus
(southernfur seals)
Phoca
Pusa
Monachini
(monk seals)
Neomonachus
Mirounga
(elephant seals)
Lobodontini
(Antarctic seals)
Ailuridae
Conepatus
(hog-nosed skunks)
Mephitis
Mydaus
(stink badgers)
Spilogale
(spotted skunks)
Bassariscus
Procyon
(raccoons)
Bassaricyon
(olingos)
Nasuina
(coatis)
Nasua
Nasuella
(mountain coatis)
Mustelidae
    • see below↓
Mellivora
Arctonyx
(hog badgers)
Meles
(Eurasian badgers)
Melogale
(ferret-badgers)
Pekania
Gulo
Martes
(martens)
Lyncodontini
Galictis
(grisons)
Ictonychini
(African polecats)
Vormela
Ictonyx
Lontra
Enhydra
Lutra
Lutrogale
Aonyx
Neogale
(New World weasels)
subgenusMustela
(paraphyletic)
subgenusLutreola
(paraphyletic)
subgenusPutorius
Game animals andshooting in the United Kingdom
Game birds
Quarry species
Deer
Other quarry
Opponents
Law
See also
1 Rarely shot because of declining numbers.
Vulpes vulpes
Canis vulpes
International
National
Other
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