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The red fox (''Vulpes vulpes'') is the largest of the true foxes and one of the most widely distributed members of the order
Carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...
, being present across the entire
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
including most of
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, plus parts of
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
. It is listed as
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. Its range has increased alongside human expansion, having been introduced to Australia, where it is considered harmful to native mammals and bird populations. Due to its presence in Australia, it is included on the list of the "world's 100 worst invasive species". The red fox originated from smaller-sized ancestors from Eurasia during the Middle
Villafranchian Villafranchian age ( ) is a period of geologic time (3.5–1.0 Ma) spanning the Late Pliocene and Early Pleistocene used more specifically with European Land Mammal Ages. Named by Italian geologist Lorenzo Pareto for a sequence of terrestrial s ...
period, and colonised North America shortly after the
Wisconsin glaciation The Wisconsin Glacial Episode, also called the Wisconsin glaciation, was the most recent glacial period of the North American ice sheet complex. This advance included the Cordilleran Ice Sheet, which nucleated in the northern North American Cor ...
. Among the true foxes, the red fox represents a more progressive form in the direction of
carnivory A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, adipose tissue, f ...
. 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 Leucism () is a wide variety of conditions that result in the partial loss of pigmentation in an animal—causing white, pale, or patchy coloration of the skin, hair, feathers, scales, or cuticles, but not the eyes. It is occasionally spelled ' ...
and melanistic individuals. Forty-five
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
are currently recognised, which are divided into two categories: the large northern foxes and the small, basal southern grey desert foxes of Asia and North Africa. Red foxes are usually found in pairs or small groups consisting of families, such as a
mated pair Breeding pair is a pair of animals which cooperate over time to produce offspring with some form of a bond between the individuals.Gaston, A. J.The evolution of group territorial behavior and cooperative breeding" The American Naturalist 112.988 ...
and their young, or a male with several females having kinship ties. The young of the mated pair remain with their parents to assist in caring for new kits. The species primarily feeds on small rodents, though it may also target
rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s,
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
s,
game bird Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are ofte ...
s,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s,
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s and young
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
s. Fruit and vegetable matter is also eaten sometimes. Although the red fox tends to kill smaller predators, including other fox species, it is vulnerable to attack from larger predators, such as
wolves The wolf (''Canis lupus''; plural, : wolves), also known as the gray wolf or grey wolf, is a large Canis, canine native to Eurasia and North America. More than thirty subspecies of Canis lupus, subspecies of ''Canis lupus'' have been reco ...
,
coyote The coyote (''Canis latrans'') is a species of canis, canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecologica ...
s,
golden jackal The golden jackal (''Canis aureus''), also called common jackal, is a wolf-like canid that is native to Southeast Europe, Southwest Asia, South Asia, and regions of Southeast Asia. The golden jackal's coat varies in color from a pale creamy y ...
s, large predatory birds such as
golden eagle The golden eagle (''Aquila chrysaetos'') is a bird of prey living in the Northern Hemisphere. It is the most widely distributed species of eagle. Like all eagles, it belongs to the family Accipitridae. They are one of the best-known bird of p ...
s and
Eurasian eagle owl The Eurasian eagle-owl (''Bubo bubo'') is a species of eagle-owl that resides in much of Eurasia. It is also called the Uhu and it is occasionally abbreviated to just the eagle-owl in Europe. It is one of the largest species of owl, and females ...
s, and medium- and large-sized felines. The species has a long history of association with humans, having been extensively hunted as a pest and
furbearer Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
for many centuries, as well as being represented in human folklore and mythology. Because of its widespread distribution and large population, the red fox is one of the most important furbearing animals harvested for the
fur trade The fur trade is a worldwide industry dealing in the acquisition and sale of animal fur. Since the establishment of a world fur market in the early modern period, furs of boreal, polar and cold temperate mammalian animals have been the mos ...
. Too small to pose a threat to humans, it has extensively benefited from the presence of human habitation, and has successfully
colonised Colonization, or colonisation, constitutes large-scale population movements wherein migrants maintain strong links with their, or their ancestors', former country – by such links, gain advantage over other inhabitants of the territory. When ...
many
suburb A suburb (more broadly suburban area) is an area within a metropolitan area, which may include commercial and mixed-use, that is primarily a residential area. A suburb can exist either as part of a larger city/urban area or as a separate ...
an and
urban Urban means "related to a city". In that sense, the term may refer to: * Urban area, geographical area distinct from rural areas * Urban culture, the culture of towns and cities Urban may also refer to: General * Urban (name), a list of people ...
areas. Domestication of the red fox is also underway in
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
, and has resulted in the
domesticated red fox The domesticated silver fox (''Vulpes vulpes'' forma ''amicus'') is a form of the silver fox that has been to some extent domesticated under laboratory conditions. The silver fox is a melanistic form of the wild red fox. Domesticated silver fox ...
.


Terminology

Males are called tods or dogs, females are called vixens, and young cubs are known as kits. Although the
Arctic fox The Arctic fox (''Vulpes lagopus''), also known as the white fox, polar fox, or snow fox, is a small fox native to the Arctic regions of the Northern Hemisphere and common throughout the Arctic tundra biome. It is well adapted to living in co ...
has a small native population in northern Scandinavia, and while the
corsac fox The corsac fox (''Vulpes corsac''), also known simply as a corsac, is a medium-sized fox found in steppes, semi-deserts and deserts in Central Asia, ranging into Mongolia and northern China. Since 2004, it has been classified as ''least concern ...
's range extends into
European Russia European Russia (russian: Европейская Россия, russian: европейская часть России, label=none) is the western and most populated part of Russia. It is geographically situated in Europe, as opposed to the cou ...
, 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
Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the early Middle Ages. It was brought to Great Britain by Anglo-Saxon settlement of Britain, Anglo ...
, which derived from
Proto-Germanic Proto-Germanic (abbreviated PGmc; also called Common Germanic) is the reconstructed proto-language of the Germanic branch of the Indo-European languages. Proto-Germanic eventually developed from pre-Proto-Germanic into three Germanic branc ...
*''fuhsaz''. Compare with West Frisian ''foks'',
Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...
''vos'', and
German German(s) may refer to: * Germany (of or related to) ** Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ge ...
''Fuchs''. This, in turn, derives from
Proto-Indo-European Proto-Indo-European (PIE) is the reconstructed common ancestor of the Indo-European language family. Its proposed features have been derived by linguistic reconstruction from documented Indo-European languages. No direct record of Proto-Indo-E ...
*''puḱ-'' 'thick-haired; tail'. Compare to the
Hindi Hindi (Devanāgarī: or , ), or more precisely Modern Standard Hindi (Devanagari: ), is an Indo-Aryan language spoken chiefly in the Hindi Belt region encompassing parts of northern, central, eastern, and western India. Hindi has been de ...
''pū̃ch'' 'tail',
Tocharian B Kuchean (also known as Tocharian B or West Tocharian) was a Western member of Tocharian branch of Indo-European languages, extinct from ninth century. Once spoken in the Tarim Basin in Central Asia. Tocharian B shows an internal chronological de ...
''päkā'' 'tail; chowrie', and Lithuanian ''paustìs'' 'fur'. The bushy tail also forms the basis for the fox's
Welsh Welsh may refer to: Related to Wales * Welsh, referring or related to Wales * Welsh language, a Brittonic Celtic language spoken in Wales * Welsh people People * Welsh (surname) * Sometimes used as a synonym for the ancient Britons (Celtic peop ...
name, ''llwynog'', literally 'bushy', from ''llwyn'' 'bush'. Likewise, pt, raposa from ''rabo'' 'tail', Lithuanian ''uodẽgis'' from ''uodegà'' 'tail', and
Ojibwe The Ojibwe, Ojibwa, Chippewa, or Saulteaux are an Anishinaabe people in what is currently southern Canada, the northern Midwestern United States, and Northern Plains. According to the U.S. census, in the United States Ojibwe people are one of ...
''waagosh'' from ''waa'', which refers to the up and down "bounce" or flickering of an animal or its tail. The scientific term ''vulpes'' derives from the Latin word for fox, and gives the adjectives ''vulpine'' and ''vulpecular''.


Evolution

The red fox is considered a more specialised form of ''Vulpes'' than the
Afghan Afghan may refer to: *Something of or related to Afghanistan, a country in Southern-Central Asia *Afghans, people or citizens of Afghanistan, typically of any ethnicity ** Afghan (ethnonym), the historic term applied strictly to people of the Pas ...
, corsac and
Bengal fox The Bengal fox (''Vulpes bengalensis''), also known as the Indian fox, is a fox endemic to the Indian subcontinent from the Himalayan foothills and Terai of Nepal through southern India, and from southern and eastern Pakistan to eastern India and ...
es in the direction of size and adaptation to
carnivory A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, adipose tissue, f ...
; the skull displays far fewer
neotenous Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compa ...
traits than in other species, and its facial area is more developed. It is, however, not as adapted for a purely carnivorous diet as the
Tibetan fox The Tibetan fox (''Vulpes ferrilata''), also known as the Tibetan sand fox, is a species of true fox endemic to the high Tibetan Plateau, Ladakh plateau, Nepal, China, Sikkim, and Bhutan, up to elevations of about . It is listed as Least Concer ...
.


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
Villafranchian Villafranchian age ( ) is a period of geologic time (3.5–1.0 Ma) spanning the Late Pliocene and Early Pleistocene used more specifically with European Land Mammal Ages. Named by Italian geologist Lorenzo Pareto for a sequence of terrestrial s ...
. The earliest fossil specimens of ''V. vulpes'' were uncovered in
Baranya Baranya or Baranja may refer to: * Baranya (region) or Baranja, a region in Hungary and Croatia * Baranya County, a county in modern Hungary * Baranya County (former), a county in the historic Kingdom of Hungary * Baranya, Hungarian name of villag ...
,
Hungary Hungary ( hu, Magyarország ) is a landlocked country in Central Europe. Spanning of the Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia a ...
dating from 3.4 to 1.8 million years ago. The ancestral species was likely smaller than the current one, as the earliest red fox fossils are smaller than modern populations. The earliest fossil remains of the modern species date back to the mid-
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
in association with the refuse of 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.


Colonisation of North America

Red foxes colonised the North American continent in two waves: before or during the
Illinoian glaciation The Illinoian Stage is the name used by Quaternary geologists in North America to designate the period c.191,000 to c.130,000 years ago, during the middle Pleistocene, when sediments comprising the Illinoian Glacial Lobe were deposited. It preced ...
, and during the Wisconsinan glaciation.
Gene mapping Gene mapping describes the methods used to identify the locus of a gene and the distances between genes. Gene mapping can also describe the distances between different sites within a gene. The essence of all genome mapping is to place a co ...
demonstrates that red foxes in North America have been isolated from their Old World counterparts for over 400,000 years, thus raising the possibility that
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
has occurred, and that the previous
binomial name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
of ''Vulpes fulva'' may be valid. In the far north, red fox fossils have been found in
Sangamonian The Sangamonian Stage (or Sangamon interglacial) is the term used in North America to designate the last interglacial period. In its most common usage, it is used for the period of time between 75,000 and 125,000 BP.Willman, H.B., and J.C. Frye, 1 ...
Stage deposits in the Fairbanks District and
Medicine Hat Medicine Hat is a city in Southern Alberta, southeast Alberta, Canada. It is located along the South Saskatchewan River. It is approximately east of Lethbridge and southeast of Calgary. This city and the adjacent Town of Redcliff, Alberta, ...
. Fossils dating from the Wisconsinan are present in 25 sites in
Arkansas Arkansas ( ) is a landlocked state in the South Central United States. It is bordered by Missouri to the north, Tennessee and Mississippi to the east, Louisiana to the south, and Texas and Oklahoma to the west. Its name is from the Osage ...
,
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
,
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...
,
Idaho Idaho ( ) is a state in the Pacific Northwest region of the Western United States. To the north, it shares a small portion of the Canada–United States border with the province of British Columbia. It borders the states of Montana and Wyom ...
,
Missouri Missouri is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. Ranking List of U.S. states and territories by area, 21st in land area, it is bordered by eight states (tied for the most with Tennessee ...
,
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
,
Tennessee Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to th ...
,
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
,
Virginia Virginia, officially the Commonwealth of Virginia, is a state in the Mid-Atlantic and Southeastern regions of the United States, between the Atlantic Coast and the Appalachian Mountains. The geography and climate of the Commonwealth ar ...
, and
Wyoming Wyoming () is a U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the south ...
. 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 American ranges because of human-induced environmental changes.
Genetic testing Genetic testing, also known as DNA testing, is used to identify changes in DNA sequence or chromosome structure. Genetic testing can also include measuring the results of genetic changes, such as RNA analysis as an output of gene expression, or ...
indicates two distinct red fox refugia 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 large subspecies ''V. v. alascensis'', ''V. v. abietorum'', ''V. v. regalis'', and ''V. v. rubricosa''. The southern (or montane) refugium occurs in the subalpine parklands and alpine meadows of the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico in ...
, the
Cascade Range The Cascade Range or Cascades is a major mountain range of western North America, extending from southern British Columbia through Washington and Oregon to Northern California. It includes both non-volcanic mountains, such as the North Cascades, ...
, and
Sierra Nevada The Sierra Nevada () is a mountain range in the Western United States, between the Central Valley of California and the Great Basin. The vast majority of the range lies in the state of California, although the Carson Range spur lies primarily ...
, and consists of the small subspecies ''V. v. cascadensis'', ''V. v. macroura'', ''V. v. necator'', and ''V. v. patwin''. The latter
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
has been separated from all other red fox populations since the last glacial maximum, and may possess unique ecological or physiological adaptations. 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 haplotypes in any North American populations. Also, introduced eastern American red foxes have colonized southern California, the
San Joaquin Valley The San Joaquin Valley ( ; es, Valle de San Joaquín) is the area of the Central Valley of the U.S. state of California that lies south of the Sacramento–San Joaquin River Delta and is drained by the San Joaquin River. It comprises seven c ...
, and
San Francisco Bay Area The San Francisco Bay Area, often referred to as simply the Bay Area, is a populous region surrounding the San Francisco, San Pablo, and Suisun Bay estuaries in Northern California. The Bay Area is defined by the Association of Bay Area Go ...
, but appear to have mixed with the
Sacramento Valley , photo =Sacramento Riverfront.jpg , photo_caption= Sacramento , map_image=Map california central valley.jpg , map_caption= The Central Valley of California , location = California, United States , coordinates = , boundaries = Sierra Nevada (ea ...
red fox (''V. v. patwin'') only in a narrow hybrid zone. In addition, no evidence is seen of interbreeding of eastern American red foxes in California with the montane
Sierra Nevada red fox The Sierra Nevada red fox (''Vulpes vulpes necator''), also known as the High Sierra fox, is a subspecies of red fox found in the Oregon Cascades and the Sierra Nevada. It is likely one of the most endangered mammals in North America. The Hig ...
(''V. v. necator'') or other populations in the Intermountain West (between the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico in ...
to the east and the Cascade and Sierra Nevada Mountains to the west).


Subspecies

The 3rd edition of ''Mammal Species of the World'' 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 A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...
haplotype A haplotype ( haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material ( DNA) which is inherited from two parents. Normally these organisms have their DNA or ...
studies. Castello (2018) recognized 30 subspecies of the Old World red fox and nine subspecies of the North American red fox as valid. Substantial
gene pool The gene pool is the set of all genes, or genetic information, in any population, usually of a particular species. Description A large gene pool indicates extensive genetic diversity, which is associated with robust populations that can surv ...
mixing between different subspecies is known; British red foxes have
crossbred A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. ''Crossbreeding'', sometimes called "designer crossbreeding", is the process of breeding such an organism, While crossbreeding is used to main ...
extensively with red foxes imported from Germany, France, Belgium, Sardinia and possibly Siberia and Scandinavia. However, genetic studies suggest very little differences between red foxes sampled across Europe. Lack of genetic diversity is consistent with the red fox being a highly agile species, with one red fox covering in under a year's time. Red fox subspecies in Eurasia and North Africa are divided into two categories: *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 Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compa ...
traits than the northern foxes 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. Red foxes living in Middle Asia show physical traits intermediate to the northern foxes and southern grey desert foxes.


Description


Build

The red fox has an elongated body and relatively short limbs. The tail, which is longer than half the body length (70 percent of head and body length), is fluffy and reaches the ground when in a standing position. Their pupils are oval and vertically oriented.
Nictitating membrane The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. All ...
s are present, but move only when the eyes are closed. The forepaws have five digits, while the hind feet have only four and lack
dewclaw A dewclaw is a digit – vestigial in some animals – on the foot of many mammals, birds, and reptiles (including some extinct orders, like certain theropods). It commonly grows higher on the leg than the rest of the foot, such that in digit ...
s. They are very agile, being capable of jumping over fences, and swim well. Vixens normally have four pairs of
teat A teat is the projection from the mammary glands of mammals from which milk flows or is ejected for the purpose of feeding young. In many mammals the teat projects from the udder. The number of teats varies by mammalian species and often corre ...
s, though vixens with seven, nine, or ten teats are not uncommon. The
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoster ...
of males are smaller than those of Arctic foxes. Their
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
s are fairly narrow and elongated, with small
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
s. Their
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
are relatively long.
Sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
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
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
s, as well as having larger canines. Their skulls are distinguished from those of dogs by their narrower muzzles, less crowded
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s, more slender canine teeth, and concave rather than convex profiles.


Dimensions

Red foxes are the largest species of the genus ''Vulpes''. However, relative to dimensions, red foxes are much lighter than similarly sized dogs of the genus ''
Canis ''Canis'' is a genus of the Caninae which includes multiple extant species, such as wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and den ...
''. Their limb bones, for example, weigh 30 percent less per unit area of bone than expected for similarly sized dogs. They display significant individual, sexual, age and geographical variation in size. On average, adults measure high at the shoulder and in body length with tails measuring . The ears measure and the hind feet . Weights range from , with vixens typically weighing 15–20% less than males.Burnie, D. & Wilson, D. E. (eds.), ''Animal: The Definitive Visual Guide to the World's Wildlife''. DK Adult (2005), Adult red foxes have skulls measuring , while those of vixens measure . The forefoot print measures in length and in width, while the hind foot print measures long and wide. They trot at a speed of , and have a maximum running speed of . They have a stride of when walking at a normal pace. 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. The largest red fox on record in Great Britain was a , long male, killed in
Aberdeenshire Aberdeenshire ( sco, Aiberdeenshire; gd, Siorrachd Obar Dheathain) is one of the 32 Subdivisions of Scotland#council areas of Scotland, council areas of Scotland. It takes its name from the County of Aberdeen which has substantially differe ...
,
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to the ...
, in early 2012.


Fur

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. Among northern foxes, the North American varieties generally have the silkiest
guard hair Guard hair or overhair is the outer layer of hair of most mammals, which overlay the fur. Guard hairs are long and coarse and protect the rest of the pelage (fur) from abrasion and frequently from moisture. They are visible on the surface of the fu ...
s, while most Eurasian red foxes have coarser fur. The fur in "thermal windows" areas such as the head and the lower legs is kept dense and short all year round, while fur in other areas changes with the seasons. The foxes actively control the peripheral
vasodilation Vasodilation is the widening of blood vessels. It results from relaxation of smooth muscle cells within the vessel walls, in particular in the large veins, large arteries, and smaller arterioles. The process is the opposite of vasoconstriction, ...
and peripheral
vasoconstriction Vasoconstriction is the narrowing of the blood vessels resulting from contraction of the muscular wall of the vessels, in particular the large arteries and small arterioles. The process is the opposite of vasodilation, the widening of blood vessel ...
in these areas to regulate heat loss. There are three main colour morphs; red, silver/black and cross (see ''
Mutations In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
''). In the typical red morph, their coats are generally bright reddish-rusty with yellowish tints. A stripe of weak, diffuse patterns of many brown-reddish-chestnut hairs occurs along the spine. Two additional stripes pass down the shoulder blades, which, 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. During lactation, the belly fur of vixens may turn brick red. The upper parts of the limbs are rusty reddish, while the paws are black. The frontal part of the face and upper neck is bright brownish-rusty 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 the supracaudal gland, is usually present at the base of the tail. The tip of the tail is white.


Colour morphs

Atypical colouration in red foxes usually represents stages toward full
melanism The term melanism refers to black pigment and is derived from the gr, μελανός. Melanism is the increased development of the dark-colored pigment melanin in the skin or hair. Pseudomelanism, also called abundism, is another variant of pi ...
, and mostly occurs in cold regions.


Senses

Red foxes have
binocular vision In biology, binocular vision is a type of vision in which an animal has two eyes capable of facing the same direction to perceive a single three-dimensional image of its surroundings. Binocular vision does not typically refer to vision where an ...
, but their sight reacts mainly to movement. Their auditory perception is acute, being able to hear
black grouse The black grouse (''Lyrurus tetrix''), also known as northern black grouse, Eurasian black grouse, blackgame or blackcock, is a large game bird in the grouse family. It is a sedentary species, spanning across the Palearctic in moorland and step ...
changing roosts at 600 paces, the flight of
crow A crow is a bird of the genus ''Corvus'', or more broadly a synonym for all of ''Corvus''. Crows are generally black in colour. The word "crow" is used as part of the common name of many species. The related term "raven" is not pinned scientifical ...
s at and the squeaking of mice at about . They are capable of locating sounds to within one degree at 700–3,000 Hz, though less accurately at higher frequencies. Their sense of smell is good, but weaker than that of specialised dogs.


Scent glands

Red foxes have a pair of
anal sac Anal may refer to: Related to the anus *Related to the anus of animals: ** Anal fin, in fish anatomy ** Anal vein, in insect anatomy ** Anal scale, in reptile anatomy *Related to the human anus: ** Anal sex, a type of sexual activity involvin ...
s lined by sebaceous glands, both of which open through a single duct. The anal sacs act as fermentation chambers in which
aerobic Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cel ...
and
anaerobic bacteria An anaerobic organism or anaerobe is any organism that does not require molecular oxygen for growth. It may react negatively or even die if free oxygen is present. In contrast, an aerobic organism (aerobe) is an organism that requires an oxygenate ...
convert
sebum A sebaceous gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in the greatest nu ...
into odorous compounds, including
aliphatic In organic chemistry, hydrocarbons ( compounds composed solely of carbon and hydrogen) are divided into two classes: aromatic compounds and aliphatic compounds (; G. ''aleiphar'', fat, oil). Aliphatic compounds can be saturated, like hexane, ...
acids. The oval-shaped
caudal gland Scent gland are exocrine glands found in most mammals. They produce semi-viscous secretions which contain pheromones and other semiochemical compounds. These odor-messengers indicate information such as status, territorial marking, mood, and Anim ...
is long and wide, and reportedly smells of
violet Violet may refer to: Common meanings * Violet (color), a spectral color with wavelengths shorter than blue * One of a list of plants known as violet, particularly: ** ''Viola'' (plant), a genus of flowering plants Places United States * Viol ...
s. 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.


Distribution and habitat

The red fox is a wide-ranging species. Its range covers nearly 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 Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland is t ...
,
Iceland Iceland ( is, Ísland; ) is a Nordic island country in the North Atlantic Ocean and in the Arctic Ocean. Iceland is the most sparsely populated country in Europe. Iceland's capital and largest city is Reykjavík, which (along with its s ...
, the Arctic islands, the most northern parts of central
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...
, and in extreme deserts. It is not present in
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
and is classed as a "prohibited new organism" under the
Hazardous Substances and New Organisms Act 1996 The Hazardous Substances and New Organisms Act (HSNO) is an Act of Parliament passed in New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North I ...
, which does not allow import.


Australia

In Australia, estimates in 2012 indicated that there were more than 7.2 million red foxes, with a range extending throughout most of the continental mainland. They became established in Australia through successive introductions in 1830s and 1840s, by settlers in the British colonies of
Van Diemen's Land Van Diemen's Land was the colonial name of the island of Tasmania used by the British during the European exploration of Australia in the 19th century. A British settlement was established in Van Diemen's Land in 1803 before it became a sepa ...
(as early as 1833) and the
Port Phillip District The Port Phillip District was an administrative division of the Colony of New South Wales from 9 September 1836 until 1 July 1851, when it was separated from New South Wales and became the Colony of Victoria. In September 1836, NSW Colonial Sec ...
of New South Wales (as early as 1845), who wanted to foster the traditional English sport of
fox hunting Fox hunting is an activity involving the tracking, chase and, if caught, the killing of a fox, traditionally a red fox, by trained foxhounds or other scent hounds. A group of unarmed followers, led by a "master of foxhounds" (or "master of ho ...
. A permanent red fox population did not establish itself on the island of
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
, and it is widely held that foxes were out-competed by the
Tasmanian devil The Tasmanian devil (''Sarcophilus harrisii'') (palawa kani: purinina) is a carnivorous marsupial of the family Dasyuridae. Until recently, it was only found on the island state of Tasmania, but it has been reintroduced to New South Wales in ...
. On the mainland, however, the species was successful as an
apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...
. The fox is generally less common in areas where the
dingo The dingo (''Canis familiaris'', ''Canis familiaris dingo'', ''Canis dingo'', or ''Canis lupus dingo'') is an ancient (Basal (phylogenetics), basal) lineage of dog found in Australia (continent), Australia. Its taxonomic classification is de ...
is more prevalent, but it has, primarily through its burrowing behaviour, achieved
niche differentiation In ecology, niche differentiation (also known as niche segregation, niche separation and niche partitioning) refers to the process by which competing species use the environment differently in a way that helps them to coexist. The competitive excl ...
with both the
feral dog A free-ranging dog is a dog that is not confined to a yard or house. Free-ranging dogs include street dogs, village dogs, stray dogs, feral dogs, etc., and may be owned or unowned. The global dog population is estimated to be 900 million, of ...
and the
feral cat A feral cat or a stray cat is an unowned domestic cat (''Felis catus'') that lives outdoors and avoids human contact: it does not allow itself to be handled or touched, and usually remains hidden from humans. Feral cats may breed over dozens ...
. Consequently, the fox has become one of the continent's most destructive invasive species. The red fox has been implicated in the extinction or decline of several native Australian species, particularly those of the family
Potoroidae Potoroidae is a family of marsupials, small Australian animals known as bettongs, potoroos, and rat-kangaroos. All are rabbit-sized, brown, jumping marsupials and resemble a large rodent or a very small wallaby. Taxonomy The potoroids are s ...
, including the
desert rat-kangaroo The desert rat-kangaroo (''Caloprymnus campestris''), also called the buff-nosed rat-kangaroo, plains rat-kangaroo or oolacunta,Tony Robinson & Tiana Forrest (2012A possible sighting of the Desert Rat-kangaroo or Oolacunta (''Caloprymnus campestr ...
. The spread of red foxes across the southern part of the continent has coincided with the spread of
rabbits in Australia European rabbits (''Oryctolagus cuniculus'') were first introduced to Australia in the 18th century with the First Fleet, and later became widespread likely because of Thomas Austin. Such wild rabbit populations are a serious mammalian pest an ...
, and corresponds with declines in the distribution of several medium-sized ground-dwelling mammals, including
brush-tailed bettong The woylie or brush-tailed bettong (''Bettongia penicillata'') is a small, critically endangered, gerbil-like mammal native to forests and shrubland of Australia. A member of the rat-kangaroo family (Potoroidae), it moves by hopping and is acti ...
s,
burrowing bettong The boodie (''Bettongia lesueur''), also known as the burrowing bettong or Lesueur's rat-kangaroo, is a small, furry, rat-like mammal native to Australia. Once common throughout the continent, it is now restricted to a few coastal islands. A memb ...
s,
rufous bettong The rufous rat-kangaroo or rufous bettong (''Aepyprymnus rufescens'') is a small, jumping, rat-like mammal native to eastern Australia. It is the only species in the genus ''Aepyprymnus''. The largest member of the rat-kangaroo family (Potoroida ...
s, bilbys,
numbat The numbat (''Myrmecobius fasciatus''), also known as the noombat or walpurti, is an insectivorous marsupial. It is diurnal and its diet consists almost exclusively of termites. The species was once widespread across southern Australia, but ...
s, bridled nail-tail wallabies and
quokka The quokka (''Setonix brachyurus'', ) is a small macropod about the size of a domestic cat. It is the only member of the genus ''Setonix''. Like other marsupials in the macropod family (such as kangaroos and wallabies), the quokka is herbivo ...
s. 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. 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 the
eastern bettong The eastern bettong (''Bettongia gaimardi''), also known as the southern or Tasmanian bettong, is a small, hopping, rat-like mammal native to grassy forests of southeastern Australia and Tasmania. A member of the rat-kangaroo family (Potoroidae) ...
, and an eradication program was initiated, conducted by the Tasmanian
Department of Primary Industries and Water The Tasmanian Department of Natural Resources and Environment (NRE) is the government department of the Tasmanian Government responsible for supporting primary industry development, the protection of Tasmania's natural environment, effective lan ...
.


Sardinia, Italy

The origin of the ''ichnusae'' subspecies in
Sardinia Sardinia ( ; it, Sardegna, label=Italian, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese and Catalan) is the second-largest island in the Mediterranean Sea, after ...
, Italy is uncertain, as it is absent from Pleistocene deposits in their current homeland. It is possible it originated during the
Neolithic The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several parts ...
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.


Behaviour


Social and territorial behaviour

Red foxes either establish stable home ranges within particular areas or are itinerant with no fixed abode. They use their urine to mark their territories. A male fox raises one hind leg and his urine is sprayed forward in front of him, whereas a female fox squats down so that the urine is sprayed in the ground between the hind legs. Urine is also used to mark empty
cache Cache, caching, or caché may refer to: Places United States * Cache, Idaho, an unincorporated community * Cache, Illinois, an unincorporated community * Cache, Oklahoma, a city in Comanche County * Cache, Utah, Cache County, Utah * Cache County ...
sites, used to store found food, as reminders not to waste time investigating them. The use of up to 12 different urination postures allows them to precisely control the position of the scent mark. Red foxes live in family groups sharing a joint territory. In favourable habitats and/or areas with low hunting pressure, subordinate foxes may be present in a range. Subordinate foxes may number one or two, sometimes up to eight in one territory. These subordinates could be formerly 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, an example of
kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like altruistic behaviour whose evolution i ...
. 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.


Reproduction and development

Red foxes reproduce once a year in spring. Two months prior to
oestrus The estrous cycle (, originally ) is the set of recurring physiological changes that are induced by reproductive hormones in most mammalian therian females. Estrous cycles start after sexual maturity in females and are interrupted by anestrous ...
(typically December), the reproductive organs of vixens change shape and size. By the time they enter their oestrus period, their
uterine horn The uterine horns (cornua of uterus) are the points in the upper uterus where the fallopian tubes exit to meet the ovaries. They are one of the points of attachment for the round ligament of uterus (the other being the mons pubis). They also provi ...
s double in size, and their
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
grow 1.5–2 times larger. Sperm formation in males begins in August–September, with the testicles attaining their greatest weight in December–February. The vixen's oestrus period lasts three weeks, during which the dog-foxes mate with the vixens for several days, often in burrows. The male's
bulbus glandis The bulbus glandis (also called a bulb or knot) is an erectile tissue structure on the penis of canid mammals. During mating, immediately before ejaculation the tissues swell up to lock ('' tie'') the male's penis inside the female. The locking i ...
enlarges during
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
, forming a
copulatory tie Canine reproduction is the process of sexual reproduction in domestic dogs, wolves, coyotes and other canine species. Canine sexual anatomy and development Male reproductive system Erectile tissue As with all mammals, a dog's penis is made up o ...
which may last for more than an hour. The
gestation period In mammals, pregnancy is the period of reproduction during which a female carries one or more live offspring from implantation in the uterus through gestation. It begins when a fertilized zygote implants in the female's uterus, and ends once it ...
lasts 49–58 days. Though foxes are largely
monogamous Monogamy ( ) is a form of Dyad (sociology), dyadic Intimate relationship, relationship in which an individual has only one Significant other, partner during their lifetime. Alternately, only one partner at any one time (Monogamy#Serial monogamy, ...
, DNA evidence from one population indicated large levels of
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any o ...
,
incest Incest ( ) is human sexual activity between family members or close relatives. This typically includes sexual activity between people in consanguinity (blood relations), and sometimes those related by affinity (marriage or stepfamily), adoption ...
and mixed paternity litters. Subordinate vixens may become pregnant, but usually fail to whelp, or have their kits killed postpartum by either the dominant female or other subordinates. The average litter size consists of four to six kits, though litters of up to 13 kits have occurred. Large litters are typical in areas where fox mortality is high. Kits are born blind, deaf and toothless, with dark brown fluffy fur. At birth, they weigh and measure in body length and in tail length. At birth, they are short-legged, large-headed and have broad chests. Mothers remain with the kits for 2–3 weeks, as they are unable to
thermoregulate Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
. During this period, the fathers or barren vixens feed the mothers. Vixens are very protective of their kits, and have been known to even fight off terriers in their defence. If the mother dies before the kits are independent, the father takes over as their provider. 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. 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. Kits begin to leave their dens and experiment with solid food brought by their parents at the age of 3–4 weeks. The
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proces ...
period lasts 6–7 weeks. Their woolly coats begin to be coated by shiny guard hairs after 8 weeks. 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. Some vixens may reach
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
at the age of 9–10 months, thus bearing their first litters at one year of age. In captivity, their
longevity The word " longevity" is sometimes used as a synonym for "life expectancy" in demography. However, the term ''longevity'' is sometimes meant to refer only to especially long-lived members of a population, whereas ''life expectancy'' is always d ...
can be as long as 15 years, though in the wild they typically do not survive past 5 years of age.


Denning behaviour

Outside the
breeding season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
, most red foxes favour living in the open, in densely vegetated areas, though they may enter burrows to escape bad weather. Their burrows are often dug on hill or mountain slopes, ravines, bluffs, steep banks of water bodies, ditches, depressions, gutters, in rock clefts and neglected human environments. Red foxes prefer to dig their burrows on well drained soils. Dens built among tree roots can last for decades, while those dug on the steppes last only several years. They may permanently abandon their dens during
mange Mange is a type of skin disease caused by parasitic mites. Because various species of mites also infect plants, birds and reptiles, the term "mange", or colloquially "the mange", suggesting poor condition of the skin and fur due to the infection ...
outbreaks, possibly as a defence mechanism against the spread of disease. In the Eurasian desert regions, foxes may use the burrows 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 are divided into a den and temporary burrows, which consist only of a small passage or cave for concealment. The main entrance of the burrow leads downwards (40–45°) and broadens into a den, from which numerous side tunnels branch. Burrow depth ranges from , rarely extending to
ground water Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidated ...
. The main passage can reach in length, standing an average of . In spring, red foxes clear their dens of excess soil through rapid movements, first with the forepaws then with kicking motions with their hind legs, throwing the discarded soil over from the burrow. When kits are born, the discarded debris is trampled, thus forming a spot where the kits can play and receive food. They may share their dens with woodchucks or badgers. Unlike badgers, which fastidiously clean their earths and defecate in
latrine A latrine is a toilet or an even simpler facility that is used as a toilet within a sanitation system. For example, it can be a communal trench in the earth in a camp to be used as emergency sanitation, a hole in the ground (pit latrine), or m ...
s, red foxes habitually leave pieces of prey around their dens. The average sleep time of a captive red fox is 9.8 hours per day.


Communication


Body language

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. 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. 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.


Vocalisations

Red foxes have a wide vocal range, and produce different sounds spanning five
octave In music, an octave ( la, octavus: eighth) or perfect octave (sometimes called the diapason) is the interval between one musical pitch and another with double its frequency. The octave relationship is a natural phenomenon that has been refer ...
s, which grade into each other. Recent analyses identify 12 different sounds produced by adults and 8 by kits. 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. *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 owl The tawny owl (''Strix aluco''), also called the brown owl, is commonly found in woodlands across Europe to western Siberia, and has seven recognized subspecies. It is a stocky, medium-sized owl, whose underparts are pale with dark streaks, an ...
s). 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. *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 Ratchet may refer to: Devices * Ratchet (device), a mechanical device that allows movement in only one direction * Ratchet, metonomic name for a socket wrench incorporating a ratcheting device * Ratchet (instrument), a music instrument and a ...
, called "gekkering". Gekkering occurs mostly during the courting season from rival males or vixens rejecting advances. Both tamed and domesticated foxes have been observed making sounds similar to laughter, which is believed to be used as a contact call when communicating with human owners and handlers. 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.


Ecology


Diet, hunting and feeding behaviour

Red foxes are
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s with a highly varied diet. Research conducted in the former Soviet Union showed red foxes consuming over 300 animal species and a few dozen species of plants. They primarily feed on small rodents like
vole Voles are small rodents that are relatives of lemmings and hamsters, but with a stouter body; a longer, hairy tail; a slightly rounder head; smaller eyes and ears; and differently formed molars (high-crowned with angular cusps instead of low-c ...
s,
mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
,
ground squirrel Ground squirrels are members of the squirrel family of rodents (Sciuridae), which generally live on or in the ground, rather than trees. The term is most often used for the medium-sized ground squirrels, as the larger ones are more commonly known ...
s,
hamster Hamsters are rodents (order Rodentia) belonging to the subfamily Cricetinae, which contains 19 species classified in seven genera.Fox, Sue. 2006. ''Hamsters''. T.F.H. Publications Inc. They have become established as popular small pets. The b ...
s,
gerbil The Mongolian gerbil or Mongolian jird (''Meriones unguiculatus'') is a small rodent belonging to the subfamily Gerbillinae. Their body size is typically , with a tail, and body weight , with adult males larger than females. The animal is use ...
s,
woodchuck The groundhog (''Marmota monax''), also known as a woodchuck, is a rodent of the family Sciuridae, belonging to the group of large ground squirrels known as marmots. The groundhog is a lowland creature of North America; it is found through mu ...
s,
pocket gopher Pocket gophers, commonly referred to simply as gophers, are burrowing rodents of the family Geomyidae. The roughly 41 speciesSearch results for "Geomyidae" on thASM Mammal Diversity Database are all endemic to North and Central America. They are ...
s and
deer mice Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...
. Secondary prey species include birds (with
Passeriformes A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
,
Galliformes Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are often ...
and
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which in ...
predominating),
leporid Leporidae is the family of rabbits and hares, containing over 60 species of extant mammals in all. The Latin word ''Leporidae'' means "those that resemble ''lepus''" (hare). Together with the pikas, the Leporidae constitute the mammalian order ...
s,
porcupine Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethizont ...
s,
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of ...
s,
opossum Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 93 species in 18 genera. Opossums originated in South America and entered North ...
s,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s,
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, other
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s,
flotsam In maritime law, flotsam'','' jetsam'','' lagan'','' and derelict are specific kinds of shipwreck. The words have specific nautical meanings, with legal consequences in the law of admiralty and marine salvage. A shipwreck is defined as the rema ...
(
marine mammals Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their re ...
,
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
and
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s) and
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
. On very rare occasions, foxes may attack young or small
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
s. They typically target mammals up to about in weight, and they require of food daily. Red foxes readily eat plant material and in some areas fruit can amount to 100% of their diet in autumn. Commonly consumed fruits include
blueberries Blueberries are a widely distributed and widespread group of perennial flowering plants with blue or purple berries. They are classified in the section ''Cyanococcus'' within the genus ''Vaccinium''. ''Vaccinium'' also includes cranberries, bi ...
,
blackberries The blackberry is an edible fruit produced by many species in the genus ''Rubus'' in the family (biology), family Rosaceae, hybrids among these species within the subgenus ''Rubus'', and hybrids between the subgenera ''Rubus'' and ''Idaeobatus' ...
,
raspberries The raspberry is the edible fruit of a multitude of plant species in the genus ''Rubus'' of the rose family, most of which are in the subgenus '' Idaeobatus''. The name also applies to these plants themselves. Raspberries are perennial with w ...
,
cherries A cherry is the fruit of many plants of the genus ''Prunus'', and is a fleshy drupe (stone fruit). Commercial cherries are obtained from cultivars of several species, such as the sweet ''Prunus avium'' and the sour ''Prunus cerasus''. The nam ...
,
persimmon The persimmon is the edible fruit of a number of species of trees in the genus ''Diospyros''. The most widely cultivated of these is the Oriental persimmon, ''Diospyros kaki'' ''Diospyros'' is in the family Ebenaceae, and a number of non-pers ...
s,
mulberries ''Morus'', a genus of flowering plants in the family Moraceae, consists of diverse species of deciduous trees commonly known as mulberries, growing wild and under cultivation in many temperate world regions. Generally, the genus has 64 identif ...
,
apple An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple fruit tree, trees are agriculture, cultivated worldwide and are the most widely grown species in the genus ''Malus''. The tree originated in Central Asia, wh ...
s,
plum A plum is a fruit of some species in ''Prunus'' subg. ''Prunus'.'' Dried plums are called prunes. History Plums may have been one of the first fruits domesticated by humans. Three of the most abundantly cultivated species are not found i ...
s,
grape A grape is a fruit, botanically a berry, of the deciduous woody vines of the flowering plant genus ''Vitis''. Grapes are a non- climacteric type of fruit, generally occurring in clusters. The cultivation of grapes began perhaps 8,000 years ago, ...
s and
acorn The acorn, or oaknut, is the nut of the oaks and their close relatives (genera ''Quercus'' and '' Lithocarpus'', in the family Fagaceae). It usually contains one seed (occasionally two seeds), enclosed in a tough, leathery shell, and borne ...
s. Other plant material includes
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...
es,
sedge The Cyperaceae are a family of graminoid (grass-like), monocotyledonous flowering plants known as sedges. The family is large, with some 5,500 known species described in about 90 genera, the largest being the "true sedges" genus ''Carex'' wit ...
s and
tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
s. Red foxes are implicated in the predation of
game A game is a structured form of play (activity), play, usually undertaken for enjoyment, entertainment or fun, and sometimes used as an educational tool. Many games are also considered to be work (such as professional players of spectator s ...
and
song bird A songbird is a bird belonging to the suborder Passeri of the perching birds (Passeriformes). Another name that is sometimes seen as the scientific or vernacular name is Oscines, from Latin ''oscen'', "songbird". The Passeriformes contains 500 ...
s, hares,
rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s,
muskrat The muskrat (''Ondatra zibethicus'') is a medium-sized semiaquatic rodent native to North America and an introduced species in parts of Europe, Asia, and South America. The muskrat is found in wetlands over a wide range of climates and habitat ...
s and young ungulates, particularly in
preserve The word preserve may refer to: Common uses * Fruit preserves, a type of sweet spread or condiment * Nature reserve, an area of importance for wildlife, flora, fauna or other special interest, usually protected Arts, entertainment, and media ...
s,
reserve Reserve or reserves may refer to: Places * Reserve, Kansas, a US city * Reserve, Louisiana, a census-designated place in St. John the Baptist Parish * Reserve, Montana, a census-designated place in Sheridan County * Reserve, New Mexico, a US vi ...
s and hunting farms where ground-nesting birds are protected and raised, as well as in
poultry farm Poultry farming is the form of animal husbandry which raises domesticated birds such as chickens, ducks, turkeys and geese to produce meat or eggs for food. Poultry – mostly chickens – are farmed in great numbers. More than 60 billion chic ...
s. While the popular consensus is that
olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
is very important for hunting, 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 and coyotes. Red foxes prefer to hunt in the early morning hours before sunrise and late evening. Although they typically forage alone, they may aggregate in resource-rich environments. 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 away. They typically only feed on carrion in the late evening hours and at night. They are extremely possessive of their food and will defend their catches from even dominant animals. Red foxes may occasionally commit acts of
surplus killing Surplus killing, also known as excessive killing, killing for sport, henhouse syndrome, or overkill, is a common behavior exhibited by predators, in which they kill more prey than they can immediately eat and then they either cache or abandon th ...
; during one breeding season, four red foxes were recorded to have killed around 200
black-headed gull The black-headed gull (''Chroicocephalus ridibundus'') is a small gull that breeds in much of the Palearctic including Europe and also in coastal eastern Canada. Most of the population is migratory and winters further south, but some birds res ...
s 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. Red foxes seem to dislike the
taste The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
of
mole Mole (or Molé) may refer to: Animals * Mole (animal) or "true mole", mammals in the family Talpidae, found in Eurasia and North America * Golden moles, southern African mammals in the family Chrysochloridae, similar to but unrelated to Talpida ...
s, but will nonetheless catch them alive and present them to their kits as playthings. A 2008–2010 study of 84 red foxes in the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
and
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
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 A magnetic field is a vector field that describes the magnetic influence on moving electric charges, electric currents, and magnetic materials. A moving charge in a magnetic field experiences a force perpendicular to its own velocity and to ...
.


Enemies and competitors

Red foxes typically dominate other fox species.
Arctic fox The Arctic fox (''Vulpes lagopus''), also known as the white fox, polar fox, or snow fox, is a small fox native to the Arctic regions of the Northern Hemisphere and common throughout the Arctic tundra biome. It is well adapted to living in co ...
es 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 The Aleutian Islands (; ; ale, Unangam Tanangin,”Land of the Aleuts", possibly from Chukchi language, Chukchi ''aliat'', "island"), also called the Aleut Islands or Aleutic Islands and known before 1867 as the Catherine Archipelago, are a cha ...
to the
Alexander Archipelago The Alexander Archipelago (russian: Архипелаг Александра) is a long archipelago (group of islands) in North America lying off the southeastern coast of Alaska. It contains about 1,100 islands, the tops of submerged coastal m ...
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. Where they are
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
, Arctic foxes may also escape competition by feeding on
lemming A lemming is a small rodent, usually found in or near the Arctic in tundra biomes. Lemmings form the subfamily Arvicolinae (also known as Microtinae) together with voles and muskrats, which form part of the superfamily Muroidea, which also include ...
s and flotsam rather than voles, as favoured by red foxes. Both species will kill each other's kits, given the opportunity. Red foxes are serious competitors of
corsac fox The corsac fox (''Vulpes corsac''), also known simply as a corsac, is a medium-sized fox found in steppes, semi-deserts and deserts in Central Asia, ranging into Mongolia and northern China. Since 2004, it has been classified as ''least concern ...
es, as they hunt the same prey all year. The red species is also stronger, is better adapted to hunting in snow deeper than 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. 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. Red foxes dominate kit and
swift fox The swift fox (''Vulpes velox'') is a small light orange-tan fox around the size of a domestic cat found in the western grasslands of North America, such as Montana, Colorado, New Mexico, Kansas, Oklahoma and Texas. It also lives in southern Ma ...
es. 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. 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.
Wolves The wolf (''Canis lupus''; plural, : wolves), also known as the gray wolf or grey wolf, is a large Canis, canine native to Eurasia and North America. More than thirty subspecies of Canis lupus, subspecies of ''Canis lupus'' have been reco ...
may kill and eat red foxes in disputes over carcasses. In areas in North America where red fox and
coyote The coyote (''Canis latrans'') is a species of canis, canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecologica ...
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. In Israel, red foxes share their habitat with
golden jackal The golden jackal (''Canis aureus''), also called common jackal, is a wolf-like canid that is native to Southeast Europe, Southwest Asia, South Asia, and regions of Southeast Asia. The golden jackal's coat varies in color from a pale creamy y ...
s. 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 In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition that two species which compete for the same limited resource cannot coexist at constant population values. When one species has even the sligh ...
. Red foxes dominate
raccoon dog The common raccoon dog (''Nyctereutes procyonoides''), also called the Chinese or Asian raccoon dog, is a small, heavy-set, fox-like canid native to East Asia. Named for its raccoon-like face markings, it is most closely related to foxes. Common ...
s, 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 The Republic of Tatarstan (russian: Республика Татарстан, Respublika Tatarstan, p=rʲɪsˈpublʲɪkə tətɐrˈstan; tt-Cyrl, Татарстан Республикасы), or simply Tatarstan (russian: Татарстан, tt ...
, 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. Red foxes may kill small
mustelid The Mustelidae (; from Latin ''mustela'', weasel) are a family of carnivorous mammals, including weasels, badgers, otters, ferrets, martens, minks and wolverines, among others. Mustelids () are a diverse group and form the largest family in ...
s like
weasel Weasels are mammals of the genus ''Mustela'' of the family Mustelidae. The genus ''Mustela'' includes the least weasels, polecats, stoats, ferrets and European mink. Members of this genus are small, active predators, with long and slender bo ...
s,
stone marten The beech marten (''Martes foina''), also known as the stone marten, house marten or white breasted marten, is a species of marten native to much of Europe and Central Asia, though it has established a feral population in North America. It is li ...
s,
pine marten The European pine marten (''Martes martes''), also known as the pine marten, is a mustelid native to and widespread in most of Europe, Asia Minor, the Caucasus and parts of Iran, Iraq and Syria. It is listed as Least Concern on the IUCN Red List. ...
s,
stoat The stoat (''Mustela erminea''), also known as the Eurasian ermine, Beringian ermine and ermine, is a mustelid native to Eurasia and the northern portions of North America. Because of its wide circumpolar distribution, it is listed as Least Conc ...
s, kolonoks,
polecat Polecat is a common name for several mustelid species in the order Carnivora and subfamilies Ictonychinae and Mustelinae. Polecats do not form a single taxonomic rank (i.e. clade). The name is applied to several species with broad similarities ...
s and young
sable The sable (''Martes zibellina'') is a species of marten, a small omnivorous mammal primarily inhabiting the forest environments of Russia, from the Ural Mountains throughout Siberia, and northern Mongolia. Its habitat also borders eastern Kaza ...
s.
Eurasian badger The European badger (''Meles meles''), also known as the Eurasian badger, is a badger species in the family Mustelidae native to almost all of Europe. It is classified as least concern on the IUCN Red List as it has a wide range and a large s ...
s may live alongside red foxes in isolated sections of large burrows. 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. However, cases are known of Eurasian badgers driving vixens from their dens and destroying their litters without eating them.
Wolverine The wolverine (), (''Gulo gulo''; ''Gulo'' is Latin for "gluttony, glutton"), also referred to as the glutton, carcajou, or quickhatch (from East Cree, ''kwiihkwahaacheew''), is the largest land-dwelling species of the family Mustelidae. It is ...
s may kill red foxes, often while the latter are sleeping or near carrion. Red foxes, in turn, may kill young wolverines. 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. Red fox remains are often found in striped hyena dens and striped hyenas may steal red foxes from traps. In Eurasia, red foxes may be preyed upon by
leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, a ...
s,
caracal The caracal (''Caracal caracal'') () is a medium-sized wild cat native to Africa, the Middle East, Central Asia, and arid areas of Pakistan and northwestern India. It is characterised by a robust build, long legs, a short face, long tufted e ...
s and
Eurasian lynx The Eurasian lynx (''Lynx lynx'') is a medium-sized wild cat widely distributed from Northern, Central and Eastern Europe to Central Asia and Siberia, the Tibetan Plateau and the Himalayas. It inhabits temperate and boreal forests up to an eleva ...
es. The Eurasian lynxes chase red foxes into deep snow, where their longer legs and larger paws give them an advantage over red foxes, especially when the depth of the snow exceeds one metre. In the Velikoluki district in Russia, red foxes are absent or are seen only occasionally where Eurasian lynxes establish permanent territories. Researchers consider Eurasian lynxes to represent considerably less danger to red foxes than wolves do. North American felid predators of red foxes include
cougar The cougar (''Puma concolor'') is a large Felidae, cat native to the Americas. Its Species distribution, range spans from the Canadian Yukon to the southern Andes in South America and is the most widespread of any large wild terrestrial mamm ...
s,
Canada lynx The Canada lynx (''Lynx canadensis''), or Canadian lynx, is a medium-sized North American lynx that ranges across Alaska, Canada, and northern areas of the contiguous United States. It is characterized by its long, dense fur, triangular ears ...
es and
bobcat The bobcat (''Lynx rufus''), also known as the red lynx, is a medium-sized cat native to North America. It ranges from southern Canada through most of the contiguous United States to Oaxaca in Mexico. It is listed as Least Concern on the IUC ...
s. Occasionally, large raptors such as
Eurasian eagle-owl The Eurasian eagle-owl (''Bubo bubo'') is a species of eagle-owl that resides in much of Palearctic, Eurasia. It is also called the Uhu and it is occasionally abbreviated to just the eagle-owl in Europe. It is one of the largest species of owl, ...
s will prey on young foxes(and adults on rare occasion), while
golden eagle The golden eagle (''Aquila chrysaetos'') is a bird of prey living in the Northern Hemisphere. It is the most widely distributed species of eagle. Like all eagles, it belongs to the family Accipitridae. They are one of the best-known bird of p ...
s,
wedge-tailed eagle The wedge-tailed eagle (''Aquila audax'') is the largest bird of prey in the continent of Australia. It is also found in southern New Guinea to the north and is distributed as far south as the state of Tasmania. Adults of this species have lon ...
s,
white-tailed eagle The white-tailed eagle (''Haliaeetus albicilla'') is a very large species of sea eagle widely distributed across temperate Palearctic, Eurasia. Like all eagles, it is a member of the family Accipitridae (or accipitrids) which includes other diur ...
s,
steller's sea eagle Steller's sea eagle (''Haliaeetus pelagicus''), also known as Pacific sea eagle or white-shouldered eagle, is a very large diurnal bird of prey in the family Accipitridae. It was described first by Peter Simon Pallas in 1811. No subspecies are r ...
s have been known to kill adults.


Diseases and parasites

Red foxes are the most important
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, vi ...
vector in Europe. In
London London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a majo ...
,
arthritis Arthritis is a term often used to mean any disorder that affects joints. Symptoms generally include joint pain and stiffness. Other symptoms may include redness, warmth, swelling, and decreased range of motion of the affected joints. In som ...
is common in foxes, being particularly frequent in the spine. Foxes may be infected with
leptospirosis Leptospirosis is a blood infection caused by the bacteria ''Leptospira''. Signs and symptoms can range from none to mild (headaches, muscle pains, and fevers) to severe ( bleeding in the lungs or meningitis). Weil's disease, the acute, severe ...
and
tularemia Tularemia, also known as rabbit fever, is an infectious disease caused by the bacterium ''Francisella tularensis''. Symptoms may include fever, skin ulcers, and enlarged lymph nodes. Occasionally, a form that results in pneumonia or a throat infe ...
, though they are not overly susceptible to the latter. They may also fall ill from
listeriosis Listeriosis is a bacterial infection most commonly caused by '' Listeria monocytogenes'', although '' L. ivanovii'' and '' L. grayi'' have been reported in certain cases. Listeriosis can cause severe illness, including severe sepsis, men ...
and spirochetosis, as well as acting as
vector Vector most often refers to: *Euclidean vector, a quantity with a magnitude and a direction *Vector (epidemiology), an agent that carries and transmits an infectious pathogen into another living organism Vector may also refer to: Mathematic ...
s in spreading
erysipelas Erysipelas () is a relatively common bacterial infection of the superficial layer of the skin ( upper dermis), extending to the superficial lymphatic vessels within the skin, characterized by a raised, well-defined, tender, bright red rash, t ...
,
brucellosis Brucellosis is a highly contagious zoonosis caused by ingestion of unpasteurized milk or undercooked meat from infected animals, or close contact with their secretions. It is also known as undulant fever, Malta fever, and Mediterranean fever. The ...
and tick-borne
encephalitis Encephalitis is inflammation of the brain. The severity can be variable with symptoms including reduction or alteration in consciousness, headache, fever, confusion, a stiff neck, and vomiting. Complications may include seizures, hallucinations, ...
. A mysterious fatal disease near
Lake Sartlan Lake Sartlan (russian: Озеро Сартлан) is an Endorheic basin, endorheic hyposaline lake in the Baraba steppe of Novosibirsk Oblast, Russia. Sartlan has a surface area of 238 km2 (92 sq mi). It is the third largest lake in Novosibir ...
in the
Novosibirsk Oblast Novosibirsk Oblast (russian: Новосиби́рская о́бласть, ''Novosibirskaya oblast'') is a federal subject of Russia (an oblast) located in southwestern Siberia. Its administrative and economic center is the city of Novosibirsk ...
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 Encephalomyelitis is inflammation of the brain and spinal cord. Various types of encephalomyelitis include: * '' Acute disseminated encephalomyelitis'' or ''postinfectious encephalomyelitis'', a demyelinating disease of the brain and spinal cord, ...
, which was first observed in captive-bred silver foxes. Individual cases of foxes infected with ''
Yersinia pestis ''Yersinia pestis'' (''Y. pestis''; formerly '' Pasteurella pestis'') is a gram-negative, non-motile, coccobacillus bacterium without spores that is related to both ''Yersinia pseudotuberculosis'' and ''Yersinia enterocolitica''. It is a facult ...
'' are known. Red foxes are not readily prone to infestation with
flea Flea, the common name for the order Siphonaptera, includes 2,500 species of small flightless insects that live as external parasites of mammals and birds. Fleas live by ingesting the blood of their hosts. Adult fleas grow to about long, a ...
s. Species like ''
Spilopsyllus cuniculi ''Spilopsyllus cuniculi'', the rabbit flea, is a species of flea in the family Pulicidae. It is an external parasite of rabbits and hares and is occasionally found on cats and dogs and also certain seabirds that nest in burrows. It can act as a ...
'' are probably only caught from the fox's prey species, while others like ''
Archaeopsylla erinacei ''Archaeopsylla'' is a genus of fleas, containing one known species, the hedgehog flea The hedgehog flea (''Archaeopsylla erinacei'') is a flea species which, as indicated by its common name, is an external parasite specifically adapted to li ...
'' are caught whilst traveling. Fleas that feed on red foxes include ''
Pulex irritans The human flea (''Pulex irritans'') – once also called the house flea – is a cosmopolitan flea species that has, in spite of the common name, a wide host spectrum. It is one of six species in the genus '' Pulex''; the other five are all confi ...
'', ''
Ctenocephalides canis The dog flea (''Ctenocephalides canis'') is a species of flea that lives as an ectoparasite on a wide variety of mammals, particularly the domestic dog and cat. It closely resembles the cat flea, ''Ctenocephalides felis'', which can live on a wid ...
'' and '' Paraceras melis''.
Tick Ticks (order Ixodida) are parasitic arachnids that are part of the mite superorder Parasitiformes. Adult ticks are approximately 3 to 5 mm in length depending on age, sex, species, and "fullness". Ticks are external parasites, living by ...
s such as ''
Ixodes ricinus ''Ixodes ricinus'', the castor bean tick, is a chiefly European species of hard-bodied tick. It may reach a length of when engorged with a blood meal, and can transmit both bacterial and viral pathogens such as the causative agents of Lyme disea ...
'' and '' I. hexagonus'' are not uncommon in red foxes, and are typically found on nursing vixens and kits still in their earths. The
louse Louse ( : lice) is the common name for any member of the clade Phthiraptera, which contains nearly 5,000 species of wingless parasitic insects. Phthiraptera has variously been recognized as an order, infraorder, or a parvorder, as a result o ...
''
Trichodectes vulpis ''Trichodectes'' is a genus of louse belonging to the family Trichodectidae. The genus was described in 1818 by Nitzsch. The genus has cosmopolitan distribution. Species: * '' Trichodectes baculus'' Schömmer, 1913 * ''Trichodectes canis ...
'' specifically targets red foxes, but is found infrequently. The
mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
''
Sarcoptes scabiei ''Sarcoptes scabiei'' or the itch mite is a parasitic mite that burrows into skin and causes scabies. The mite is found in all parts of the world. Humans are not the only mammals that can become infected. Other mammals, such as wild and domesti ...
'' is the most important cause of
mange Mange is a type of skin disease caused by parasitic mites. Because various species of mites also infect plants, birds and reptiles, the term "mange", or colloquially "the mange", suggesting poor condition of the skin and fur due to the infection ...
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 the
epizootic In epizoology, an epizootic (from Greek: ''epi-'' upon + ''zoon'' animal) is a disease event in a nonhuman animal population analogous to an epidemic in humans. An epizootic may be restricted to a specific locale (an "outbreak"), general (an "epi ...
phase of the disease, it usually takes red foxes four months to die after infection. Other endoparasites include ''
Demodex folliculorum ''Demodex folliculorum'' is a microscopic mite that can only survive on the skin of humans. Most people have ''D.folliculorum'' on their skin. Usually, the mites do not cause any harm, so are considered an example of commensalism rather than par ...
'', '' Notoderes'', ''
Otodectes cynotis ''Otodectes'' is a genus of parasitic mite in the superfamily Psoroptidae (the biting cohort of the Sarcoptiformes order of mites). There is a single species of mite in this genus, ''Otodectes cynotis''. This mite is within the class Arachnida an ...
'' (which is frequently found in the
ear canal The ear canal (external acoustic meatus, external auditory meatus, EAM) is a pathway running from the outer ear to the middle ear. The adult human ear canal extends from the pinna to the eardrum and is about in length and in diameter. Struc ...
), ''
Linguatula serrata ''Linguatula serrata'' is a species of cosmopolitan zoonotic parasite, belonging to the tongueworm order Pentastomida. They are wormlike parasites of the respiratory systems of vertebrates. They live in the nasopharyngeal region of mammals. Cat ...
'' (which infects the nasal passages) and
ringworm Dermatophytosis, also known as ringworm, is a fungal infection of the skin. Typically it results in a red, itchy, scaly, circular rash. Hair loss may occur in the area affected. Symptoms begin four to fourteen days after exposure. Multiple a ...
s. Up to 60
helminth Parasitic worms, also known as helminths, are large macroparasites; adults can generally be seen with the naked eye. Many are intestinal worms that are soil-transmitted and infect the gastrointestinal tract. Other parasitic worms such as schi ...
species are known to infect captive-bred foxes in
fur farm Fur farming is the practice of breeding or raising certain types of animals for their fur. Most of the world's farmed fur is produced by European farmers. In 2018, there were 5,000 fur farms in the EU, all located across 22 countries; these ...
s, while 20 are known in the wild. Several
coccidia Coccidia (Coccidiasina) are a subclass of microscopic, spore-forming, single-celled obligate intracellular parasites belonging to the apicomplexan class Conoidasida. As obligate intracellular parasites, they must live and reproduce within an a ...
n species of the
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
''
Isospora ''Isospora'' is a genus of internal parasites in the subclass Coccidia. It is responsible for the condition isosporiasis, which causes acute, non-bloody diarrhoea in immunocompromised individuals. Taxonomy At least 248 species were originall ...
'' and ''
Eimeria ''Eimeria'' is a genus of apicomplexan parasites that includes various species capable of causing the disease coccidiosis in animals such as cattle, poultry and smaller ruminants including sheep and goats. ''Eimeria'' species are considered to be ...
'' are also known to infect them. The most common
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
species found in red fox guts are ''
Toxocara canis ''Toxocara canis'' (also known as dog roundworm) is a worldwide-distributed helminth parasite of dogs and other canids. The name is derived from the Greek word "," meaning bow or quiver, and the Latin word "caro," meaning flesh.Bassert , J., & T ...
'' and ''
Uncinaria stenocephala ''Uncinaria stenocephala'' is a nematode that parasitizes dogs, cats, and foxes as well as humans. It is rare to find in cats in the United States. ''Uncinaria stenocephala'' is the most common canine hookworm in cooler regions, such as Canada ...
'', ''
Capillaria aerophila ''Capillaria aerophila'' is a nematode parasite found in the respiratory tract of foxes, dogs, and various other carnivorous mammals. A few cases of human infestation have also been reported. Though it is sometimes called a "lungworm", this t ...
'' and ''
Crenosoma vulpis ''Crenosoma'' is a genus of nematodes belonging to the family Crenosomatidae. The species of this genus are found in Europe and Northern America. Species: * ''Crenosoma brasiliense'' Vieira, Muniz-Pereira, Lima, Moraes Neto, Guimaraes & Luque, ...
''; the latter two infect their lungs. ''
Capillaria plica ''Capillaria plica'' (dog bladder worm) is a parasitic nematode which is most often found in the urinary bladder, and occasionally in the kidneys, of dogs and foxes. It has also been found in the domestic cat, and various wild mammals. Its ...
'' infects the red fox's bladder. ''
Trichinella spiralis ''Trichinella spiralis'' is a viviparous nematode parasite, occurring in rodents, pigs, bears, hyenas and humans, and is responsible for the disease trichinosis. It is sometimes referred to as the "pork worm" due to it being typically encounte ...
'' rarely affects them. The most common
tapeworm Eucestoda, commonly referred to as tapeworms, is the larger of the two subclasses of flatworms in the class Cestoda (the other subclass is Cestodaria). Larvae have six posterior hooks on the scolex (head), in contrast to the ten-hooked Cestodar ...
species in red foxes are '' Taenia spiralis'' and '' T. pisiformis''. Others include ''
Echinococcus granulosus ''Echinococcus granulosus,'' also called the hydatid worm, hyper tape-worm or dog tapeworm, is a cyclophyllid cestode that dwells in the small intestine of canids as an adult, but which has important intermediate hosts such as livestock and hu ...
'' and '' E. multilocularis''. Eleven
trematode Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive host ...
species infect red foxes, including ''
Metorchis conjunctus ''Metorchis conjunctus'', common name Canadian liver fluke, is a species of trematode parasite in the family Opisthorchiidae. It can infect mammals that eat raw fish in North America. The first intermediate host is a freshwater snail and the ...
''.


Relationships with humans


In folklore, religion and mythology

Red foxes feature prominently in the folklore and mythology of human cultures with which they are sympatric. In
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the Cosmogony, origin and Cosmology#Metaphysical co ...
, the
Teumessian fox In Greek mythology, the Teumessian fox, or Cadmean vixen, was a gigantic fox that was destined never to be caught. (''Teumēs(s)íā alôpēx''), ''gen''.: Τευμησίας ἀλώπεκος, also known as ἀλώπηξ τῆς Τευμησσο ...
, or Cadmean vixen, was a gigantic fox that was destined never to be caught. The fox was one of the children of
Echidna Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the ...
. In
Celtic mythology Celtic mythology is the body of myths belonging to the Celtic peoples.Cunliffe, Barry, (1997) ''The Ancient Celts''. Oxford, Oxford University Press , pp. 183 (religion), 202, 204–8. Like other Iron Age Europeans, Celtic peoples followed a ...
, the red fox is a symbolic animal. In the
Cotswolds The Cotswolds (, ) is a region in central-southwest England, along a range of rolling hills that rise from the meadows of the upper Thames to an escarpment above the Severn Valley and Evesham Vale. The area is defined by the bedrock of Jur ...
,
witch Witchcraft traditionally means the use of Magic (supernatural), magic or supernatural powers to harm others. A practitioner is a witch. In Middle Ages, medieval and early modern Europe, where the term originated, accused witches were usually ...
es were thought to take the shape of foxes to steal
butter Butter is a dairy product made from the fat and protein components of churned cream. It is a semi-solid emulsion at room temperature, consisting of approximately 80% butterfat. It is used at room temperature as a spread, melted as a condiment ...
from their neighbours. In later European folklore, the figure of
Reynard the Fox Reynard the Fox is a literary cycle of medieval allegorical Dutch, English, French and German fables. The first extant versions of the cycle date from the second half of the 12th century. The genre was popular throughout the Late Middle Ages, a ...
symbolises trickery and deceit. He originally appeared (then under the name of "Reinardus") as a secondary character in the 1150 poem "
Ysengrimus ''Ysengrimus'' is a Latin fabliau and mock epic, an anthropomorphic series of fables written in 1148 or 1149, possibly by the poet Nivardus. Its chief character is Isengrim, the Wolf; the plot describes how the trickster figure Reynard, the Fo ...
". He reappeared in 1175 in Pierre Saint Cloud's ''Le Roman de Renart'', and made his debut in England in
Geoffrey Chaucer Geoffrey Chaucer (; – 25 October 1400) was an English poet, author, and civil servant best known for ''The Canterbury Tales''. He has been called the "father of English literature", or, alternatively, the "father of English poetry". He wa ...
's ''
The Nun's Priest's Tale "The Nun's Priest's Tale" (Middle English: ''The Nonnes Preestes Tale of the Cok and Hen, Chauntecleer and Pertelote'') is one of '' The Canterbury Tales'' by the Middle English poet Geoffrey Chaucer. Composed in the 1390s, it is a beast fabl ...
''. 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. Chinese folk tales tell of fox-spirits called ''
huli jing Huli jing () are Chinese mythological creatures usually capable of shapeshifting, who may either be benevolent or malevolent spirits. In Chinese mythology and folklore, the fox spirit takes variant forms with different meanings, powers, charac ...
'' that may have up to nine tails, or ''
kumiho A kumiho or gumiho (, literally " nine-tailed fox") is a creature that appears in the folktales on East Asia and legends of Korea. It is similar to the Chinese and the Japanese . It can freely transform, among other things, into a beautiful woma ...
'' as they are known in Korea. In
Japanese mythology Japanese mythology is a collection of traditional stories, folktales, and beliefs that emerged in the islands of the Japanese archipelago. Shinto and Buddhist traditions are the cornerstones of Japanese mythology. The history of thousands of year ...
, the
kitsune In Japanese folklore, , are foxes that possess paranormal abilities that increase as they get older and wiser. According to ''yōkai'' folklore, all foxes have the ability to shapeshift into human form. While some folktales speak of employing t ...
are fox-like spirits possessing magical abilities that increase with their age and wisdom. Foremost among these is the ability to assume human form. While some folktales speak of kitsune employing this ability to trick others, other stories portray them as faithful guardians, friends, lovers, and wives. In
Arab folklore Arab culture is the culture of the Arabs, from the Atlantic Ocean in the west to the Arabian Sea in the east, and from the Mediterranean Sea in the north to the Horn of Africa and the Indian Ocean in the southeast. The various religions the Arab ...
, 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. The animal's cunning was noted by the authors of the
Bible The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity, Judaism, Samaritanism, and many other religions. The Bible is an anthologya compilation of texts of a ...
who applied the word "fox" to false prophets (
Ezekiel Ezekiel (; he, יְחֶזְקֵאל ''Yəḥezqēʾl'' ; in the Septuagint written in grc-koi, Ἰεζεκιήλ ) is the central protagonist of the Book of Ezekiel in the Hebrew Bible. In Judaism, Christianity, and Islam, Ezekiel is acknow ...
13:4) and the hypocrisy of
Herod Antipas Herod Antipas ( el, Ἡρῴδης Ἀντίπας, ''Hērǭdēs Antipas''; born before 20 BC – died after 39 AD), was a 1st-century ruler of Galilee and Perea, who bore the title of tetrarch ("ruler of a quarter") and is referred to as both "H ...
(
Luke People *Luke (given name), a masculine given name (including a list of people and characters with the name) *Luke (surname) (including a list of people and characters with the name) *Luke the Evangelist, author of the Gospel of Luke. Also known as ...
13:32). The cunning Fox is commonly found in
Native American mythology The indigenous peoples of the Americas comprise numerous different cultures. Each has its own mythologies. Some are quite distinct, but certain themes are shared across the cultural boundaries. Northern America There is no single mythology o ...
, where it is portrayed as an almost constant companion to
Coyote The coyote (''Canis latrans'') is a species of canis, canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecologica ...
. Fox, however, is a deceitful companion that often steals Coyote's food. In the
Achomawi Achomawi (also Achumawi, Ajumawi and Ahjumawi), are the northerly nine (out of eleven) bands of the Pit River tribe of Palaihnihan Native Americans who live in what is now northeastern California in the United States. These 5 autonomous ban ...
creation myth A creation myth (or cosmogonic myth) is a symbolic narrative of how the world began and how people first came to inhabit it., "Creation myths are symbolic stories describing how the universe and its inhabitants came to be. Creation myths develop ...
, Fox and Coyote are the co-creators of the world, that leave just before the arrival of humans. The
Yurok tribe The Yurok (Karuk language: Yurúkvaarar / Yuru Kyara - "downriver Indian; i.e. Yurok Indian") are an Indigenous people from along the Klamath River and Pacific coast, whose homelands are located in present-day California stretching from Trinidad ...
believed that Fox, in anger, captured the
Sun The Sun is the star at the center of the Solar System. It is a nearly perfect ball of hot plasma, heated to incandescence by nuclear fusion reactions in its core. The Sun radiates this energy mainly as light, ultraviolet, and infrared radi ...
, and tied him to a hill, causing him to burn a great hole in the ground. An
Inuit Inuit (; iu, ᐃᓄᐃᑦ 'the people', singular: Inuk, , dual: Inuuk, ) are a group of culturally similar indigenous peoples inhabiting the Arctic and subarctic regions of Greenland, Labrador, Quebec, Nunavut, the Northwest Territories ...
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 The Menominee (; mez, omǣqnomenēwak meaning ''"Menominee People"'', also spelled Menomini, derived from the Ojibwe language word for "Wild Rice People"; known as ''Mamaceqtaw'', "the people", in the Menominee language) are a federally recog ...
story tells of how Fox is an untrustworthy friend to Wolf.


Hunting

The earliest historical records of fox hunting come from the 4th century BC;
Alexander the Great Alexander III of Macedon ( grc, wikt:Ἀλέξανδρος, Ἀλέξανδρος, Alexandros; 20/21 July 356 BC – 10/11 June 323 BC), commonly known as Alexander the Great, was a king of the Ancient Greece, ancient Greek kingdom of Maced ...
is known to have hunted foxes and a
seal Seal may refer to any of the following: Common uses * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impr ...
dated from 350 BC depicts a
Persian Persian may refer to: * People and things from Iran, historically called ''Persia'' in the English language ** Persians, the majority ethnic group in Iran, not to be conflated with the Iranic peoples ** Persian language, an Iranian language of the ...
horseman in the process of spearing a fox.
Xenophon Xenophon of Athens (; grc, wikt:Ξενοφῶν, Ξενοφῶν ; – probably 355 or 354 BC) was a Greek military leader, philosopher, and historian, born in Athens. At the age of 30, Xenophon was elected commander of one of the biggest Anci ...
, who viewed hunting as part of a cultured man's education, advocated the killing of foxes as pests, as they distracted hounds from hares. The
Romans Roman or Romans most often refers to: *Rome, the capital city of Italy * Ancient Rome, Roman civilization from 8th century BC to 5th century AD *Roman people, the people of ancient Rome *''Epistle to the Romans'', shortened to ''Romans'', a lette ...
were hunting foxes by AD 80. During the Dark Ages in Europe, foxes were considered secondary quarries, but gradually grew in importance.
Cnut the Great Cnut (; ang, Cnut cyning; non, Knútr inn ríki ; or , no, Knut den mektige, sv, Knut den Store. died 12 November 1035), also known as Cnut the Great and Canute, was King of England from 1016, King of Denmark from 1018, and King of Norwa ...
re-classed foxes as Beasts of the Chase, 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 Edward I (17/18 June 1239 – 7 July 1307), also known as Edward Longshanks and the Hammer of the Scots, was King of England and Lord of Ireland from 1272 to 1307. Concurrently, he ruled the duchies of Aquitaine and Gascony as a vassal o ...
had a royal pack 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 his ''
The Master of Game ''The Master of Game'' is a medieval manuscript, a hunting treatise, written by Edward of Norwich, 2nd Duke of York, between 1406 and 1413, of which 27 manuscripts survive. It is considered to be the oldest English-language book on hunting. ''Th ...
''. By the
Renaissance The Renaissance ( , ) , from , with the same meanings. is a period in European history marking the transition from the Middle Ages to modernity and covering the 15th and 16th centuries, characterized by an effort to revive and surpass ideas ...
, fox hunting became a traditional sport of the nobility. After the
English Civil War The English Civil War (1642–1651) was a series of civil wars and political machinations between Parliamentarians (" Roundheads") and Royalists led by Charles I ("Cavaliers"), mainly over the manner of England's governance and issues of re ...
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. Although already native to North America, red foxes from England were imported for sporting purposes to Virginia and
Maryland Maryland ( ) is a state in the Mid-Atlantic region of the United States. It shares borders with Virginia, West Virginia, and the District of Columbia to its south and west; Pennsylvania to its north; and Delaware and the Atlantic Ocean to ...
in 1730 by prosperous tobacco planters. These American fox hunters considered the red fox more sporting than the grey fox. 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 Saskatchewan ( ; ) is a Provinces and territories of Canada, province in Western Canada, western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on t ...
(Canada) 2,000 (2000–2001);
Nova Scotia Nova Scotia ( ; ; ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
(Canada) 491 (2000–2001);
Minnesota Minnesota () is a state in the upper midwestern region of the United States. It is the 12th largest U.S. state in area and the 22nd most populous, with over 5.75 million residents. Minnesota is home to western prairies, now given over to ...
(US) 4,000–8,000 (average annual trapping harvest 2002–2009);
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
(US) 69 (1999–2000).


Fur use

Red foxes are among the most important fur-bearing animals harvested by the
fur trade The fur trade is a worldwide industry dealing in the acquisition and sale of animal fur. Since the establishment of a world fur market in the early modern period, furs of boreal, polar and cold temperate mammalian animals have been the mos ...
. Their pelts are used for
trim Trim or TRIM may refer to: Cutting * Cutting or trimming small pieces off something to remove them ** Book trimming, a stage of the publishing process ** Pruning, trimming as a form of pruning often used on trees Decoration * Trim (sewing), or ...
mings,
scarf A scarf, plural ''scarves'', is a piece of fabric worn around the neck or head for warmth, sun protection, cleanliness, fashion, religious reasons, or used to show the support for a sports club or team. They can be made in a variety of diff ...
s, muffs,
jacket A jacket is a garment for the upper body, usually extending below the hips. A jacket typically has sleeves, and fastens in the front or slightly on the side. A jacket is generally lighter, tighter-fitting, and less insulating than a coat, which ...
s and
coat A coat typically is an outer garment for the upper body as worn by either gender for warmth or fashion. Coats typically have long sleeves and are open down the front and closing by means of buttons, zippers, hook-and-loop fasteners, toggles, a ...
s. They are principally used as trimming for both cloth coats and fur garments, including evening wraps. The pelts of silver foxes are popular as
cape A cape is a clothing accessory or a sleeveless outer garment which drapes the wearer's back, arms, and chest, and connects at the neck. History Capes were common in medieval Europe, especially when combined with a hood in the chaperon. Th ...
s, while cross foxes are mostly used for scarves and rarely for trimming. 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. The silver colour morphs are the most valued by furriers, followed by the cross colour morphs and the red colour morphs, respectively. 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. 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. Pelt prices are increasing, with 2012 North American wholesale auction prices averaging $39 and 2013 prices averaging $65.78. 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. 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.


Livestock and pet predation

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 The Blackface or Scottish Blackface is a British breed of sheep. It is the most common sheep breed of the United Kingdom. Despite the name, it did not originate in Scotland, but south of the border. History The origins of the breed are ...
, are more vulnerable than larger breeds, such as the
Merino The Merino is a breed or group of breeds of domestic sheep, characterised by very fine soft wool. It was established in Spain near the end of the Middle Ages, and was for several centuries kept as a strict Spanish monopoly; exports of the bree ...
. 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. Red foxes may prey on
domestic rabbit A domestic or domesticated rabbit (''Oryctolagus cuniculus domesticus'')—more commonly known as a pet rabbit, bunny, bun, or bunny rabbit—is a subspecies of European rabbit, a member of the lagomorph family. A male rabbit is known as a ''bu ...
s and
guinea pig The guinea pig or domestic guinea pig (''Cavia porcellus''), also known as the cavy or domestic cavy (), is a species of rodent belonging to the genus ''Cavia'' in the family Caviidae. Breeders tend to use the word ''cavy'' to describe the ani ...
s 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
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s 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.


Taming and domestication

In their unmodified wild state, red foxes are generally unsuitable as pets. 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. 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. 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. They maintain their wild counterparts' strong instinct of concealment and may pose a threat to domestic birds, even when well-fed. 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. 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 of
domesticated silver fox The domesticated silver fox (''Vulpes vulpes'' forma ''amicus'') is a form of the silver fox that has been to some extent domesticated under laboratory conditions. The silver fox is a melanistic form of the wild red fox. Domesticated silver f ...
es was developed by
Russian Russian(s) refers to anything related to Russia, including: *Russians (, ''russkiye''), an ethnic group of the East Slavic peoples, primarily living in Russia and neighboring countries *Rossiyane (), Russian language term for all citizens and peo ...
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 A piebald or pied animal is one that has a pattern of unpigmented spots (white) on a pigmented background of hair, feathers or scales. Thus a piebald black and white dog is a black dog with white spots. The animal's skin under the white backgro ...
coats, floppy ears in kits and curled tails, similar to the traits that distinguish domestic dogs from grey wolves.


Urban red foxes


Distribution

Red foxes have been exceedingly successful in colonising built-up environments, especially lower-density suburbs, although many have also been sighted in dense 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, entering
Bristol Bristol () is a city, ceremonial county and unitary authority in England. Situated on the River Avon, it is bordered by the ceremonial counties of Gloucestershire to the north and Somerset to the south. Bristol is the most populous city in ...
and London during the 1940s, and later established themselves in
Cambridge Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
and
Norwich Norwich () is a cathedral city and district of Norfolk, England, of which it is the county town. Norwich is by the River Wensum, about north-east of London, north of Ipswich and east of Peterborough. As the seat of the See of Norwich, with ...
. In Ireland, they are now common in suburban
Dublin Dublin (; , or ) is the capital and largest city of Republic of Ireland, Ireland. On a bay at the mouth of the River Liffey, it is in the Provinces of Ireland, province of Leinster, bordered on the south by the Dublin Mountains, a part of th ...
. In Australia, red foxes were recorded in
Melbourne Melbourne ( ; Boonwurrung/Woiwurrung: ''Narrm'' or ''Naarm'') is the capital and most populous city of the Australian state of Victoria, and the second-most populous city in both Australia and Oceania. Its name generally refers to a met ...
as early as the 1930s, while in Zurich, Switzerland, they only started appearing in the 1980s. Urban red foxes are most common in residential suburbs consisting of privately owned, low-density housing. They are rare in areas where industry,
commerce Commerce is the large-scale organized system of activities, functions, procedures and institutions directly and indirectly related to the exchange (buying and selling) of goods and services among two or more parties within local, regional, nation ...
or council-rented houses predominate. 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 , whereas those in more residential areas average from . In 2006, it was estimated that there were 10,000 red foxes in London. 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.


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.


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. 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. 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.


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
University of Brighton The University of Brighton is a public university based on four campuses in Brighton and Eastbourne on the south coast of England. Its roots can be traced back to 1858 when the Brighton School of Art was opened in the Royal Pavilion. It achieve ...
in partnership with the BBC TV series ''
Winterwatch ''Springwatch'', ''Autumnwatch'' and ''Winterwatch'', sometimes known collectively as ''The Watches'', are annual BBC television series which chart the fortunes of British wildlife during the changing of the seasons in the United Kingdom. The p ...
'', had unexpectedly traveled 195 miles in 21 days from his neighbourhood in
Hove Hove is a seaside resort and one of the two main parts of the city of Brighton and Hove, along with Brighton in East Sussex, England. Originally a "small but ancient fishing village" surrounded by open farmland, it grew rapidly in the 19th cen ...
at the western edge of
East Sussex East Sussex is a ceremonial and non-metropolitan county in South East England on the English Channel coast. It is bordered by Kent to the north and east, West Sussex to the west, and Surrey to the north-west. The largest settlement in East Su ...
across rural countryside as far as Rye, at the eastern edge of the county. He was still continuing his journey when the
GPS The Global Positioning System (GPS), originally Navstar GPS, is a Radionavigation-satellite service, satellite-based radionavigation system owned by the United States government and operated by the United States Space Force. It is one of t ...
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. File:Red fox crossing road.jpg, An urban red fox crossing a street File:Flickr - Duncan~ - Fox Trot.jpg, An urban red fox in central London File:Fox with food bag.jpg, An urban red fox eating from a bag of biscuits in Dorset, England File:Fleet Fox AKA Rolf, Winterwatch Brighton.png, "Fleet", the urban red fox from the BBC TV series ''
Winterwatch ''Springwatch'', ''Autumnwatch'' and ''Winterwatch'', sometimes known collectively as ''The Watches'', are annual BBC television series which chart the fortunes of British wildlife during the changing of the seasons in the United Kingdom. The p ...
'' File:Red Fox eating KFC.webm, An urban red fox with a discarded
KFC KFC (Kentucky Fried Chicken) is an American fast food restaurant chain headquartered in Louisville, Kentucky, that specializes in fried chicken. It is the world's second-largest restaurant chain (as measured by sales) after McDonald's, with 2 ...
bag


References


Further reading

* * *


External links

*
Red Fox
National Geographic
Natural History of the Red Fox
Wildlife Online
Sacramento Valley red fox info1Red Fox
Fletcher Wildlife Garden {{Good article Vulpes Holarctic fauna Carnivorans of Asia Carnivorans of Europe Carnivorans of North America Mammals of the Arctic Arctic land animals Introduced mammals of Australia Mammals described in 1758 Taxa named by Carl Linnaeus Articles containing video clips Symbols of Mississippi Extant Middle Pleistocene first appearances