Vulpes vulpes (Linnaeus, 1758) Frisch, 1775

Don E. Wilson & Russell A. Mittermeier, 2009, Canidae, Handbook of the Mammals of the World – Volume 1 Carnivores, Barcelona: Lynx Edicions, pp. 352-446 : 441-442

27.

Red Fox

Vulpes vulpes

French: Renard roux / German: Rotfuchs / Spanish: Zorro rojo

Other common names: Silver Fox, Cross Fox

Taxonomy. Canis vulpes Linnaeus, 1758 ,

Sweden.

The North American Red Fox, V. fulva , previously considered a separate species, is now considered conspecific with the Palaearctic V. vulpes . Forty-four subspecies were described on the basis of regional variation, but these have doubtful ecological significance as evidenced by successful introductions and re-introductions around the world.

Subspecies and Distribution.

V. v. vulpes Linnaeus, 1758 — N Europe (Scandinavia).

V. v. abietorum Merriam, 1900 — SW Canada (Alberta & British Columbia).

V. v. aegyptiacus Sonnini, 1816 — Egypt, Israel, and Lybia.

V. v. alascensis Merriam, 1900 — Alaska and NW Canada (NW Territories & Yukon).

V. v. alpheraky: Satunin, 1906 — Kazakhstan.

V. v. anatolica Thomas, 1920 — Turkey.

V. v. arabica Thomas, 1902 — Arabian peninsula.

V. v. atlantica Wagner, 1841 — Algeria (forested Atlas Mts).

V. v. bangsi Merriam, 1900 — NE Canada (Labrador).

V. v. barbara Shaw, 1800 — NW Africa (Barbary Coast).

V. v. beringiana Middendorff, 1875 — NE Siberia (shore of Bering Strait).

V. v. cascadensis Merriam, 1900 — NW USA (Cascade Mountains, Oregon & Washington).

V. v. caucasica Dinnik, 1914 — SW Russia (Caucasus).

V. v. crucigera Bechstein, 1789 — Europe through N & C Russia.

V. v. daurica Ognev, 1931 — E Russia (Amur, Siberia & Transbaikalia).

V.v. deletrix Bangs, 1898 — NE Canada (Newfoundland).

V. v. dolichocrania Ognev, 1926 — SE Siberia (S Ussuri).

V. v. flavescens Gray, 1843 — N Iran.

V. v. fulva Desmarest, 1820 — E USA.

V. v. griffith: Blyth, 1854 — Afghanistan and N Pakistan.

V.v. harrimani Merriam, 1900 — Alaska (Kodiak I).

V. v. hoole Swinhoe, 1870 — S China (Fujian to Sichuan).

V. v. ichnusae G. S. Miller, 1907 — Corsica and Sardinia.

V. v. induta G. S. Miller, 1907 — Cyprus.

V. v. jakutensis Ognev, 1923 — E Siberia (S of Yakutsk).

V. v. japonica Gray, 1868 — Japan.

V. v. karagan Erxleben, 1777 — Mongolia, Kazakhstan, and Kirgizstan.

V. v. kenaiensis Merriam, 1900 — Alaska (Kenai Peninsula).

V. v. kurdistanica Satunin, 1906 — Armenia and NE Turkey.

V. v. macroura Baird, 1852 — USA (Mountain States).

V. v. montana Pearson, 1836 — Himalayas form China (Yunnan) to C Pakistan.

V. v. mecator Merriam, 1900 — SW USA (California & Nevada).

V. v. ochroxantha Ognev, 1926 — E Russian Turkestan, Aksai, Kirgizstan, Semirechie.

V. v. palaestina Thomas, 1920 — Jordan and Lebanon.

V.v. peculiosa Kishida, 1924 — Korea.

V. v. pusilla Blyth, 1854 — NW India to Irak.

V.v. regalis Merriam, 1900 — N Great Plains of Canada and USA.

V. v. rubricosa Bangs, 1898 — E Canada.

V.v. schrencki Kishida, 1924 — N Japan (Hokkaido) and NE Russia (Sakhalin).

V. v. silacea G. S. Miller, 1907 — Iberian Peninsula.

V.v. splendidissima Kishida, 1924 — E Russia (N & C Kurile Is).

V. v. stepensis Brauner, 1914 — steppes of S Russia.

V. v. tobolica Ognev, 1926 — Russia (lower basin of Ob River)

V. v. tschiliensis Matschie, 1907 — NE China.

Foxes of European origin were introduced into E USA and Canada in the 17" century, subsequently mixed with local subspecies. Also introduced to Australia in 1800s, and the Falkland Islands ( Malvinas).

Descriptive notes. Head-body 59-90 cm for males and 45-68 cm for females, tail 36-44 cm for males and 28-49 cm for females; weight 4-14 kg for males and 3-7 kg for females. The largest fox in the genus Vulpes , with an enormous geographical variation in size. Males generally larger than females. The species is substantially smaller in the Middle East deserts than in Europe. Smaller also in North America. Muzzle slender and pointed, white on upper lip. Ears large, pointed, erect and black-backed. Pelage is reddish-brown but may vary from brown to russet red to yellowish-gray. Three main color morphs: red,silver, and cross (grayish-brown with long black guard hairs down back and across shoulders). Throat and/or chest may have white markings. Legs long and slender. Lower legs black. Tail long, thick, and bushy, sometimes with white tip. Red Foxes in North America are comparatively light weight, rather long for their mass, and with considerable sexual dimorphism. British foxes are heavier but relatively short. European foxes are closer to the general average among populations. Body mass and length are positively related to latitude (i.e. Bergmann’s Rule), but this is a smaller effect than that related to geographical origin. Dental formula 13/3, C 1/1, PM 3/4, M 3/3 = 42.

Habitat. Red Foxes recorded in habitats as diverse as tundra, desert, grassland, and forest, as well as in cities. Natural habitat is dry, mixed landscape, with mosaics of scrub, woodland and farmland. They are also abundant on moorlands, mountains, deserts, sand dunes and farmland from sea level to 4500 m. Red Foxes flourish in urban areas, where they are most common in low-density residential suburbs and less common in areas dominated by industry or commerce. In many habitats, foxes appear to be closely associated with humans, even thriving in intensive agricultural areas. Also in large cities, including Washington DC, London, Paris, and Stockholm.

Food and Feeding. Adaptable and opportunistic omnivores, with a diet ranging from invertebrates (e.g. earthworms and beetles) to mammals, birds, fruit, and carrion. As predators, foxes typically kill birds and mammals up to about 3-5 kg. They cache food that is in excess of their requirements and have a highly developed memory for locations of hoards. They scavenge in rural areas (e.g. on deer and sheep carcasses in Europe and Canada, which may be the major food source in upland areas in winter), and in urban areas (on bird-feeding tables, compost heaps, and refuse). They are independent and thus generally solitary foragers, although individuals may forage in close proximity where resources are clumped.

Activity patterns. Foraging is mainly nocturnal and crepuscular, although more diurnal where undisturbed.

Movements, Home range and Social organization. The basic social unit is a pair, but groups with up to six members (usually one adult male and 2-5 probably related females) may share a territory, depending on habitat. Range size is habitat-dependent and can vary from less than 0-4 km? to over 40 km ®. There are reports of overlapping home ranges in some urban and rural environments, and of drifting territories in other urban settings. Mating behavior is highly variable, either monogamous pairs, a single male with two breeding females that may or may not share a communal den, or a single breeding female with several non-breeding female helpers. There is always only one breeding male in the group, although additional matings do occur outside the group. Territorial males make frequent excursions beyond their territories during the mating season, during which itinerant males also make incursions into territories. There is socially-mediated suppression of reproduction amongst females, with lowest productivity tending to occur where fox density is high or food supply poor. Where food is not limited, social status itself can suppress reproduction, with only the dominant female breeding. Behavioral mechanisms by which this occurs include harassment of subordinates, infanticide and cannibalism of subordinate vixens’ pups. A hormonal mechanism whereby stress leads to lowered productivity through fetal reabsorption has also been identified. In populations where productivity is low, reproductive performance is suppressed consistently at all stages of pregnancy from conception to birth. Juveniles may disperse between six and twelve months of age, mostly between October and January. All or most males disperse, but the proportion of each sex dispersing varies among habitats and may depend on extent of mortality (e.g. due to rabies or control). Males typically disperse farther than females. Dispersal distances generally range from less than 5 km to more than 50 km, but distances of up to 394 km have been recorded in the USA. Red Foxes communicate with facial expressions, vocalizations, and scent marking. Scent marking involves urine and feces, anal sac secretions, the violet or supracaudal gland (more active in males during breeding season), and glands around thelips, in the angle of the jaw, and between pads of the feet. Some 28 different categories of vocalization have been described, used to communicate over long distances and at close quarters.

Breeding. Males are seasonally fecund. Mating occurs between December and February (June to October in Australia); the onset of breeding is correlated with day length and so starts earlier at more southerly latitudes. Following a gestation period of 49-55 days, births occur from March to May. Lactation lasts for four weeks, and the pups are fully weaned at 6-8 weeks. Sexual maturity is reached at 9-10 months. The proportion of breeding females in the group, andlitter size (3-12 young perlitter), varies with food availability. Fox populations that are dense relative to food resources are generally less productive than those that are less dense. A single litter per year is the norm. In high-density populations where interactions with the dominant female are high, subordinate females do not usually breed. The male provides food to the lactating female, who is generally confined to the den prior to weaning. Food is provided for the pups by both parents. Non-breeding females may also feed, groom, and tend pups and have been known to adopt orphaned pups. If two females breed within a group, they may share a den and litters may be communally suckled.

Status and Conservation. CITES notlisted. Classified as Least Concern on The IUCN Red List. Widely regarded as a pest and unprotected. V. v. necator in the Sierra Nevada, California ( USA) is rare, possibly declining. The subspecies griffithi, montana, and pusilla (= leucopus) are listed as CITES Appendix III ( India). Most countries and/or states where trapping or hunting occurs have regulated closed versus open seasons and restrictions on methods of capture. In the EU, Canada, and the Russian Federation, trapping methods are regulated. Red Fox density is highly variable, ranging within the UK from 0-025 individuals/km? in Scotland to 30/km? in some urban areas where food is superabundant. Density in mountainous rural areas of Switzerland is 3/km?. In northern boreal forests and Arctic tundra, the foxes occur at densities of 0-1/km?, and in southern Ontario, Canada at 1/km? Threats include habitat degradation and fragmentation, exploitation and persecution, and government policies. Other threats come from local, national, or international socio-economic and political factors, such as increasing human population and thus increasing development. The Red fox’s versatility and eclectic diet are likely to ensure its persistence despite changes in landscape and prey base. There are currently bounties in Pakistan to protect game birds such as houbara bustards (Chlamydotis undulata macqueenii).

Bibliography. Ables (1975), Baker & Harris (2004), Cavallini (1995), Doncaster & Macdonald (1991), Englund (1970), Harris (1977, 1989), Harris & Rayner (1986), Harris & Smith (1987), Hatting (1956), Heydon & Reynolds (2000), Johnson & Hersteinsson (1993), Lariviere & Pasitschniak-Arts (1996), Macdonald (1976, 1977a, 1977b, 1979b, 1980a, 1980b, 1987), Macdonald & Bacon (1982), Macdonald & Newdick (1982), Macdonald & Reynolds (2004), Macdonald et al. (1999), McIntosh (1963), Meia (1994), Meia & Weber (1996), Voigt & Macdonald (1984).

Figures

On following pages: 28. Corsac Fox (Vulpes corsac); 29. Tibetan Fox (Vulpes ferrilata); 30. Indian Fox (Vulpes bengalensis); 31. Pale Fox (Vulpes pallida); 32. Riippell's Fox (Vulpes rueppellii; 33. Cape Fox (Vulpes chamal); 34. Blanford’s Fox (Vulpes cana); 35. Fennec Fox (Vulpes zerda).

Subspecies and Distribution. V. v. vulpes Linnaeus, 1758 — N Europe (Scandinavia). V. v. abietorum Merriam, 1900 — SW Canada (Alberta & British Columbia). V. v. aegyptiacus Sonnini, 1816 — Egypt , Israel , and Lybia. V. v. alascensis Merriam, 1900 — Alaska and NW Canada (NW Territories & Yukon). V. v. alpheraky: Satunin, 1906 — Kazakhstan . V. v. anatolica Thomas, 1920 — Turkey . V. v. arabica Thomas, 1902 — Arabian peninsula. V. v. atlantica Wagner, 1841 — Algeria (forested Atlas Mts). V. v. bangsi Merriam, 1900 — NE Canada (Labrador). V. v. barbara Shaw, 1800 — NW Africa (Barbary Coast). V. v. beringiana Middendorff, 1875 — NE Siberia (shore of Bering Strait). V. v. cascadensis Merriam, 1900 — NW USA (Cascade Mountains, Oregon & Washington). V. v. caucasica Dinnik, 1914 — SW Russia (Caucasus). V. v. crucigera Bechstein, 1789 — Europe through N & C Russia . V. v. daurica Ognev, 1931 — E Russia (Amur, Siberia & Transbaikalia). V.v. deletrix Bangs, 1898 — NE Canada (Newfoundland). V. v. dolichocrania Ognev, 1926 — SE Siberia (S Ussuri). V. v. flavescens Gray, 1843 — N Iran . V. v. fulva Desmarest, 1820 — E USA . V. v. griffith: Blyth, 1854 — Afghanistan and N Pakistan . V.v. harrimani Merriam, 1900 — Alaska (Kodiak I). V. v. hoole Swinhoe, 1870 — S China (Fujian to Sichuan). V. v. ichnusae G. S. Miller, 1907 — Corsica and Sardinia. V. v. induta G. S. Miller, 1907 — Cyprus . V. v. jakutensis Ognev, 1923 — E Siberia (S of Yakutsk). V. v. japonica Gray, 1868 — Japan . V. v. karagan Erxleben, 1777 — Mongolia , Kazakhstan , and Kirgizstan. V. v. kenaiensis Merriam, 1900 — Alaska (Kenai Peninsula). V. v. kurdistanica Satunin, 1906 — Armenia and NE Turkey . V. v. macroura Baird, 1852 — USA (Mountain States). V. v. montana Pearson, 1836 — Himalayas form China (Yunnan) to C Pakistan . V. v. mecator Merriam, 1900 — SW USA (California & Nevada). V. v. ochroxantha Ognev, 1926 — E Russian Turkestan, Aksai, Kirgizstan, Semirechie. V. v. palaestina Thomas, 1920 — Jordan and Lebanon . V.v. peculiosa Kishida, 1924 — Korea . V. v. pusilla Blyth, 1854 — NW India to Irak . V.v. regalis Merriam, 1900 — N Great Plains of Canada and USA . V. v. rubricosa Bangs, 1898 — E Canada . V.v. schrencki Kishida, 1924 — N Japan (Hokkaido) and NE Russia (Sakhalin). V. v. silacea G. S. Miller, 1907 — Iberian Peninsula. V.v. splendidissima Kishida, 1924 — E Russia (N & C Kurile Is). V. v. stepensis Brauner, 1914 — steppes of S Russia . V. v. tobolica Ognev, 1926 — Russia (lower basin of Ob River) V. v. tschiliensis Matschie, 1907 — NE China . Foxes of European origin were introduced into E USA and Canada in the 17" century, subsequently mixed with local subspecies. Also introduced to Australia in 1800s, and the Falkland Islands ( Malvinas ).