Thestoat (Mustela erminea), also known as theEurasian ermine orermine, is a species ofmustelid native toEurasia and the northern regions ofNorth America. Because of its widecircumpolar distribution, it is listed asLeast Concern on theIUCN Red List.[1]The name ermine (/ˈɜːrmɪn/) is used especially in its pure white winter coat of the stoat or its fur.[2] Ermine fur was used in the 15th century byCatholic monarchs, who sometimes used it as themozzetta cape. It has long been used on the ceremonial robes of members of the United KingdomHouse of Lords. It was also used in capes on images such as theInfant Jesus of Prague.
The stoat wasintroduced into New Zealand in the late 19th century to control rabbits. However, they have had a devastating effect on native bird populations; as such, the species was nominated as one of the world's top 100 "worst invaders".[3]
Theroot word for "stoat" is likely either the Dutch wordstout ("bold")[4] or theGothic word𐍃𐍄𐌰𐌿𐍄𐌰𐌽 (stautan, "to push").[5] According toJohn Guillim, in hisDisplay of Heraldrie, the word "ermine" is likely derived from Armenia, the nation where it was thought the species originated,[4] though other authors have linked it to theNorman French from theTeutonicharmin (Anglo-Saxonhearma). This seems to come from theLithuanian wordšarmu.[5] In North America it is called a short-tailed weasel. A male stoat is called adog,hob, orjack, while a female is called ajill. Thecollective noun for stoats is eithergang orpack.[6]
Formerly considered a single species with a very wide circumpolar range, a 2021 study splitM. erminea into three species:M. ermineasensu stricto (Eurasia and northern North America),M. richardsonii (most of North America), andM. haidarum (several islands off thePacific Northwest coast).[7][8][9]
alpestris (Burg, 1920) giganteus (Burg, 1920) major (Nilsson, 1820)
Tundra stoatM. e. arctica
Merriam, 1826
A large subspecies, with a dark yellowish-brown summer coat, a deep yellow underbelly and a massive skull; it resembles the Eurasian stoat subspecies more closely than any other American stoat subspecies[11]
A small subspecies; it has a very light, straw-brownish or greyish coat, which is short and soft. Light spots, sometimes forming a collar, are present on the neck. It does not turn white in winter.[12][13]
Tien Shan andPamir-Alay mountains, Afghanistan, India, western Tibet and the adjacent parts of the Tien Shan in China
shnitnikovi (Ognev, 1935)
whiteheadi (Wroughton, 1908)
Irish stoatM. e. hibernica
Thomas and Barrett-Hamilton, 1895
Larger thanaestiva, but smaller thanstabilis. It is distinguished by the irregular pattern on the dividing line between the dark and pale fur on the flanks, though 13.5% of Irish stoats exhibit the more typical straight dividing line.[14] They do not turn white in winter.[15][16]
East Siberian stoat (known locally as Ezo stoat in Japan) M. e. kaneii
Baird, 1857
A moderately sized subspecies. It is smaller thanM. e. tobolica, with close similarities toM. e. arctica. The colour of the summer coat is relatively light, with varying intensities of browning-yellow tinges.[17]
A moderately sized subspecies with less dense fur thanM. e. tobolica. The colour of its summer coat consists of weakly developed reddish-brown tones. The skull is similar to that ofM. e. aestiva.[17]
The mountains of southern Siberia eastwards toBaikal and the contiguous parts of Mongolia
The stoat's direct ancestor wasMustela palerminea, a common carnivore in central and eastern Europe during theMiddle Pleistocene,[20] that spread to North America during the lateBlancan or earlyIrvingtonian.[21] The stoat is the product of a process that began 5–7 million years ago, when northern forests were replaced by open grassland, thus prompting an explosiveevolution of small, burrowing rodents. The stoat's ancestors were larger than the current form, and underwent a reduction in size as they exploited the new food source. The stoat first arose in Eurasia, shortly after thelong-tailed weasel, which is in a different genus (Neogale), arose as its mirror image in North America2 million years ago. The stoat thrived during theIce Age, as its small size and long body allowed it to easily operate beneath snow, as well as hunt in burrows. The stoat and the long-tailed weasel remained separated until 500,000 years ago, when fallingsea levels exposed theBering land bridge.[22]
Fossilised stoat remains have been recovered fromDenisova Cave.[23] Combinedphylogenetic analyses indicate the stoat's closest living relatives are theAmerican ermine (M. richardsonii) andHaida ermine (M. haidarum), the latter of which partially descends fromM. erminea.[7] It is basal to most other members ofMustela, with only theyellow-bellied (M. kathia),Malayan (M. katiah), andback-striped (M. strigidorsa) weasels being more basal.[24] Themountain weasel (Mustela altaica) was formerly considered its closest relative although more recent analyses have found it to be significantly more derived. It was also previously thought to be allied with members of the genusNeogale such as the long-tailed weasel, but as those species have since been separated into a new genus, this is likely not the case.[25]
Stoat (left) and least weasel (right) pelts—note the stoat's larger size and black tail-tip
The stoat is similar to the least weasel in general proportions, manner of posture, and movement, though the tail is relatively longer, always exceeding a third of the body length,[clarification needed][26] though it is shorter than that of the long-tailed weasel. The stoat has an elongated neck, the head being set exceptionally far in front of the shoulders. The trunk is nearly cylindrical, and does not bulge at the abdomen. The greatest circumference of body is littlemore than half its length.[27] The skull, although very similar to that of the least weasel, is relatively longer, with a narrowerbraincase. The projections of the skull and teeth are weakly developed, but stronger than those of the least weasel.[28] The eyes are round, black and protrude slightly. Thewhiskers are brown or white in colour, and very long. The ears are short, rounded and lie almost flattened against the skull. The claws are not retractable, and are large in proportion to the digits. Each foot has five toes. The male stoat has a curvedbaculum with a proximal knob that increases in weight as it ages.[29]Fat is deposited primarily along the spine and kidneys, then on gut mesenteries, under the limbs and around the shoulders. The stoat has four pairs of nipples, though they are visible only in females.[29]
Skeleton
The dimensions of the stoat are variable, but not as significantly as the least weasel's.[30] Unusual among the Carnivora, the size of stoats tends to decrease proportionally with latitude, in contradiction toBergmann's rule.[20]Sexual dimorphism in size is pronounced, with males being roughly 25% larger than females and 1.5-2.0 times their weight.[14] On average, males measure 187–325 mm (7.4–12.8 in) in body length, while females measure 170–270 mm (6.7–10.6 in). The tail measures 75–120 mm (3.0–4.7 in) in males and 65–106 mm (2.6–4.2 in) in females. In males, the hind foot measures 40.0–48.2 mm (1.57–1.90 in), while in females it is 37.0–47.6 mm (1.46–1.87 in). The height of the ear measures 18.0–23.2 mm (0.71–0.91 in) in males and 14.0–23.3 mm (0.55–0.92 in). The skulls of males measure 39.3–52.2 mm (1.55–2.06 in) in length, while those of females measure 35.7–45.8 mm (1.41–1.80 in). Males average 258 g (9.1 oz) in weight, while females weigh less than 180 g (6.3 oz).[30]
The stoat has large analscent glands measuring 8.5 mm × 5 mm (0.33 in × 0.20 in) in males and smaller in females. Scent glands are also present on the cheeks, belly and flanks.[29]Epidermal secretions, which are deposited duringbody rubbing, are chemically distinct from the products of the anal scent glands, which contain a higher proportion of volatile chemicals. When attacked or being aggressive, the stoat secretes the contents of itsanal glands, giving rise to a strong, musky odour produced by several sulphuric compounds. The odour is distinct from that of least weasels.[31]
The winter fur is very dense and silky, but quite closely lying and short, while the summer fur is rougher, shorter and sparse.[26] In summer, the fur is sandy-brown on the back and head and a white below. The division between the dark back and the light belly is usually straight, though this trait is only present in 13.5% of Irish stoats. The stoat moults twice a year. In spring, the moult is slow, starting from the forehead, across the back, toward the belly. In autumn, the moult is quicker, progressing in the reverse direction. The moult, initiated byphotoperiod, starts earlier in autumn and later in spring at higher latitudes. In the stoat's northern range, it adopts a completely white coat (save for the black tail-tip) during the winter period.[29] Differences in the winter and summer coats are less apparent in southern forms of the species.[32] In the species' southern range, the coat remains brown, but is denser and sometimes paler than in summer.[29]
The stoat has acircumboreal range throughout North America, Europe, and Asia. The stoat in Europe is found as far south as 41ºN in Portugal, and inhabits most islands with the exception of Iceland,Svalbard, theMediterranean islands and some smallNorth Atlantic islands. In Japan, it is present in central mountains (northern and centralJapanese Alps) to northern part ofHonshu (primarily above 1,200 m) and Hokkaido. Its vertical range is from sea level to 3,000 m (9,800 ft).[1] In North America, it is found throughoutAlaska and westernYukon to most ofArctic Canada east toGreenland. Throughout the rest of North America, as well as parts ofNunavut, includingBaffin Island and some islands in southeast Alaska, it is replaced byM. richardsonii.[7]
Stoats have been present inOrkney, north of Scotland, since 2010, where they are a predator of theOrkney vole[33][34] and native bird populations.[35] In 2018, a stoat eradication plan, the Orkney Native Wildlife Project, was applied across the archipelago.[36][37]By 2024 the Orkney Native Wildlife Project had spent £7.9m trapping more than 6,300 stoats.[38]
Stoats were introduced intoNew Zealand during the late 19th century to control rabbits and hares, but are now a major threat to native bird populations. The introduction of stoats was opposed by scientists in New Zealand and Britain, including the New Zealand ornithologistWalter Buller. The warnings were ignored and stoats began to be introduced from Britain in the 1880s, resulting in a noticeable decline in bird populations within six years.[39] Stoats are a serious threat to ground- and hole-nesting birds, since the latter have very few means of escaping predation. The highest rates of stoat predation occur after seasonal gluts in southernbeechmast (beechnuts), which enable the reproduction of rodents on which stoats also feed, enabling stoats to increase their own numbers.[40] For instance, the endangeredSouth Island takahē's wild population dropped by a third between 2006 and 2007, after a stoat plague triggered by the 2005–06mast wiped out more than half the takahē in untrapped areas.[41]
In the Northern Hemisphere, mating occurs in the April–July period. In spring, the male'stestes are enlarged, a process accompanied by an increase oftestosterone concentration in theplasma.Spermatogenesis occurs in December, and the males are fertile from May to August, after which the testes regress.[42] Female stoats are usually onlyin heat for a brief period, which is triggered by changes in day length.[43]Copulation can last as long as 1 hour.[44] Stoats are notmonogamous, with litters often being of mixed paternity. Stoats undergoembryonic diapause, meaning that the embryo does not immediately implant in the uterus after fertilization, but rather lies dormant for a period of nine to ten months.[45] Thegestation period is therefore variable but typically around 300 days, and after mating in the summer, the offspring will not be born until the following spring – adult female stoats spend almost all their lives either pregnant or in heat.[43] Females can reabsorb embryos and in the event of a severe winter they may reabsorb their entire litter.[46] Males play no part in rearing the young, which are born blind, deaf, toothless and covered in fine white or pinkish down. Themilk teeth erupt after three weeks, and solid food is eaten after four weeks. The eyes open after five to six weeks, with the black tail-tip appearing a week later.Lactation ends after 12 weeks. Prior to the age of five to seven weeks, kits have poorthermoregulation, so they huddle for warmth when the mother is absent. Males become sexually mature at 10–11 months, while females are sexually mature at the age of 2–3 weeks whilst still blind, deaf and hairless, and are usually mated with adult males before beingweaned.[47]
Stoat territoriality has a generally mustelid spacing pattern, with male territories encompassing smaller female territories, which they defend from other males. The size of the territory and the ranging behaviour of its occupants varies seasonally, depending on the abundance of food and mates. During the breeding season, the ranges of females remain unchanged, while males either become roamers, strayers or transients. Dominant older males have territories 50 times larger than those of younger, socially inferior males. Both sexesmark their territories withurine,feces and two types ofscent marks; anal drags are meant to convey territorial occupancy, and body rubbing is associated with agonistic encounters.[31]
The stoat does not dig its own burrows, instead using the burrows and nest chambers of the rodents it kills. The skins and underfur of rodent prey are used to line the nest chamber. The nest chamber is sometimes located in seemingly unsuitable places, such as among logs piled against the walls of houses. The stoat also inhabits old and rotting stumps, under tree roots, in heaps of brushwood, haystacks, in bog hummocks, in the cracks of vacant mud buildings, in rock piles, rock clefts, and even inmagpie nests. Males and females typically live apart, but close to each other.[48] Each stoat has several dens dispersed within its range. A single den has several galleries, mainly within 30 cm (12 in) of the surface.[49]
As with theleast weasel, mouse-like rodents predominate in the stoat's diet. It regularly preys on larger rodent andlagomorph species, and takes individuals far larger than itself. In Russia, its prey includes rodents and lagomorphs such asEuropean water voles,common hamsters,pikas and others, which it overpowers in their burrows. Prey species of secondary importance include smallbirds,fish, andshrews and, more rarely,amphibians,lizards, andinsects.[50] It also preys onlemmings.[51]In Great Britain,European rabbits are an important food source, with the frequency in which stoats prey on them having increased between the 1960s and mid 1990s since the end of themyxomatosis epidemic. Typically, male stoats prey on rabbits more frequently than females do, which depend to a greater extent on smaller rodent species. British stoats rarely kill shrews,rats,squirrels and water voles, though rats may be an important food source locally. In Ireland, shrews and rats are frequently eaten. In mainland Europe, water voles make up a large portion of the stoat's diet.Hares are sometimes taken, but are usually young specimens.[52] In New Zealand, the stoat feeds principally on birds, including the rarekiwi,kaka,mohua,yellow-crowned parakeet, andNew Zealand dotterel.[52] Cases are known of stoats preying on youngmuskrats. The stoat typically eats about 50 g (1.8 oz) of food a day, which is equivalent to 25% of the animal's live weight.[53]
The stoat is an opportunistic predator that moves rapidly and checks every available burrow or crevice for food. Because of their larger size, male stoats are less successful than females in pursuing rodents far into tunnels. Stoats regularly climb trees to gain access to birds' nests, and are common raiders of nest boxes, particularly those of large species. The stoat reputedly mesmerises prey such as rabbits by a "dance" (sometimes called theweasel war dance), though this behaviour could be linked toSkrjabingylus infections.[52] The stoat seeks to immobilize large prey such as rabbits with a bite to the spine at the back of the neck. The stoat maysurplus kill when the opportunity arises; excess prey is usuallycached and eaten later. Overweight stoats would be at a disadvantage when pursuing prey into their burrows.[54] Small prey typically die instantly from a bite to the back of the neck, while larger prey, such as rabbits, typically die ofshock, as the stoat's canine teeth are too short to reach the spinal column or major arteries.[52]
The stoat is a usually silent animal; however, it can produce a range of sounds similar to those of the least weasel. Kits produce a fine chirping noise. Adults trill excitedly before mating, and indicate submission through quiet trilling, whining and squealing. When nervous, the stoat hisses, and will intersperse this with sharp barks or shrieks and prolonged screeching when aggressive.[31]
Aggressive behavior in stoats is categorized in these forms:[31]
Noncontact approach, which is sometimes accompanied by a threat display and vocalization from the approached animal
Forward thrust, accompanied by a sharp shriek, which is usually done by stoats defending a nest or retreat site
Nest occupation, when a stoat appropriates the nesting site of a weaker individual
Kleptoparasitism, in which a dominant stoat appropriates the kill of a weaker one, usually after a fight.
Submissive stoats express their status by avoiding higher-ranking animals, fleeing from them or making whining or squealing sounds.[31]
Tuberculosis has been recorded in stoats inhabiting the former Soviet Union and New Zealand. They are largely resistant totularemia, but are reputed to suffer fromcanine distemper in captivity. Symptoms ofmange have also been recorded.[58]
Stoats are vulnerable toectoparasites associated with their prey and the nests of other animals on which they do not prey. ThelouseTrichodectes erminea is recorded in stoats living in Canada, Ireland and New Zealand. In continental Europe, 26flea species are recorded to infest stoats, includingRhadinospylla pentacantha,Megabothris rectangulatus,Orchopeas howardi,Spilopsyllus ciniculus,Ctenophthalamus nobilis,Dasypsyllus gallinulae,Nosopsyllus fasciatus,Leptospylla segnis,Ceratophyllus gallinae,Parapsyllus n. nestoris,Amphipsylla kuznetzovi andCtenopsyllus bidentatus.Tick species known to infest stoats areIxodes canisuga,I. hexagonus,I. ricinus andHaemaphysalis longicornis. Louse species known to infest stoats includeMysidea picae andPolyplax spinulosa.Mite species known to infest stoats includeNeotrombicula autumnalis,Demodex erminae,Eulaelaps stabulans,Gymnolaelaps annectans,Hypoaspis nidicorva, andListrophorus mustelae.[58]
InIrish mythology, stoats were viewedanthropomorphically as animals with families, which held rituals for their dead. They were also viewed as noxious animals prone to thieving, and their saliva was said to be able to poison a grown man. To encounter a stoat when setting out for a journey was considered bad luck, but one could avert this by greeting the stoat as a neighbour.[59] Stoats were also supposed to hold the souls of infants who died beforebaptism.[60]
In the folklore of theKomi people of theUrals, stoats are symbolic of beautiful and coveted young women.[61] In theZoroastrian religion, the stoat is considered a sacred animal, as its white winter coat represented purity. Similarly,Mary Magdalene was depicted as wearing a white stoat pelt as a sign of her reformed character.
One popular European legend had it that a white stoat would die before allowing its pure white coat to be besmirched. When it was being chased by hunters, it would supposedly turn around and give itself up to the hunters rather than risk soiling itself.[62]
Stoat skins are prized by the fur trade, especially in winter coat, and used to trim coats and stoles. The fur from the winter coat is referred to as ermine and is an ancient symbol of theDuchy of Brittany, forming its earliest flag. There is also a design calledermine inspired by the winter coat of the stoat and painted onto other furs, such as rabbit.[63] In Europe these furs are a symbol of royalty and high status. Theceremonial robes of members of the United Kingdompeerage and theacademic hoods of the universities of Oxford andCambridge are traditionally trimmed with ermine.[63] In practice, rabbit or fake fur is now often used due to expense oranimal rights concerns.Prelates of the Catholic Church still wear ecclesiastical garments featuring ermine (a sign of their status equal to that of the nobility).Cecilia Gallerani is depicted holding an ermine in her portrait,Lady with an Ermine, byLeonardo da Vinci.Henry Peacham'sEmblem 75, which depicts an ermine being pursued by a hunter and two hounds, is entitledCui candor morte redemptus ("Purity Bought with His Own Death"); Peacham goes on to preach that men and women should follow the example of the ermine and keep their minds and consciences as pure as the legendary ermine keeps its fur.[64]
Ermine inheraldry is afur, a type oftincture, consisting of a white background with a pattern of black shapes representing the winter coat of the stoat. The linings of medieval coronation cloaks and some other garments, usually reserved for use by high-ranking peers and royalty, were made by sewing many ermine furs together to produce a luxurious white fur with patterns of hanging black-tipped tails. Due largely to the association of the ermine fur with the linings of coronation cloaks, crowns and peerage caps, the heraldic tincture of ermine was usually reserved to similar applications in heraldry (i.e., the linings ofcrowns andchapeaux and of theroyal canopy).[65]
Ermine (bothM. erminea andM. richardsonii, both of which inhabited the Tlingit's territory) were also valued by theTlingit and otherindigenous peoples of the Pacific Northwest Coast. They could be attached to traditional regalia and cedar bark hats as status symbols, or they were also made into shirts.[66]
The stoat was a fundamental item in the fur trade ofRussia until the 20th century. At times, no less than half the global catch came from within the borders of the former Soviet Union, containing the highest grades of stoat pelts. Stoat harvesting never became a specialty in any part of Russia, with most stoats being captured either in box-traps or jaw-traps, or with dogs. Dogs have typically been used to capture stoats incidentally near villages in Russia, and less often as part of pre-planned hunts. Guns were rarely used, as they could damage the pelt.[67]
Three Mighty Ladies from Livonia byAlbrecht Dürer (1521) watercolour, Paris, Musee duLouvre
American actress Alice Maison shown wearing ermine fur in aMack Sennett comedy film
Thea Sternheim, wife of playwrightCarl Sternheim, wearing an ermine hat
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