Least weasel

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The least weasel (Mustela nivalis), little weasel, common weasel, or simply weasel is the smallest member of the genus Mustela, family Mustelidae and order Carnivora. It is native to Eurasia, North America and North Africa, and has been introduced to New Zealand, Malta, Crete, the Azores, and São Tomé. It is classified as least concern by the IUCN, due to its wide distribution and large population throughout the Northern Hemisphere.<ref name=iucn/>

The least weasel varies greatly in size over its range. The body is slender and elongated, and the legs and tail are relatively short. The colour varies geographically, as does the pelage type and length of tail. The dorsal surface, flanks, limbs and tail of the animal are usually some shade of brown while the underparts are white. The line delineating the boundary between the two colours is usually straight. At high altitudes and in the northern part of its range, the coat becomes pure white in winter. Eighteen subspecies are recognised.

Small rodents form the largest part of the least weasel's diet, but it also kills and eats rabbits, other mammals, and occasionally birds, birds' eggs, fish and frogs. Males mark their territories with olfactory signals and have exclusive home ranges which may intersect with or include several female ranges. Least weasels use pre-existing holes to sleep, store food and raise their young. Breeding takes place in the spring and summer, and there is a single litter of about six kits which are reared exclusively by the female. Due to its small size and fierce nature, the least weasel plays an important part in the mythology and legend of various cultures.

Taxonomy and evolutionEdit

The least weasel was given its scientific name Mustela nivalis by Carl Linnaeus in his 12th edition of Systema Naturae in 1766; its epithet nivalis comes from the Latin word nix meaning "snow" because it grows a white coat during the winter season.<ref name=MS>Template:Cite journal</ref>Template:Rp The type locality was Västerbotten in Sweden.<ref name=MS/>Template:Rp As an animal with a very wide distribution, the morphology of the least weasel varies geographically. The species was reviewed by Reichstein in 1957 and again by van Zyll de Jong in 1992 and Reig in 1997. Youngman (1982) placed it in the subgenus Mustela while Abramov (1999) considered it should be included in the subgenus Gale. Based on skull characteristics, Reig (1997) proposed that the taxon should be split into four species, M. subpalmata, M. rixosa, M. vulgaris and M. eskimo. Abrimov and Baryshinikov (2000) disagreed, recognising only M. subpalmata (the Egyptian weasel) as a separate species.<ref>Template:Cite book</ref> However, Rodrigues et al. (2016) recognized M. subpalmata as a distinct population of nivalis rather than a distinct species.<ref>Template:Cite journal</ref>

Within the genus Mustela, the least weasel is a relatively unspecialised form, as evidenced by its pedomorphic skull, which occurs even in large subspecies.<ref name=s972>Template:Harvnb</ref> Its direct ancestor was Mustela praenivalis, which lived in Europe during the Middle Pleistocene and Villafranchian. M. praenivalis itself was probably preceded by M. pliocaenica of the Pliocene. The modern species probably arose during the Late Pleistocene.<ref name=k102>Template:Harvnb</ref> The least weasel is the product of a process begun 5–7 million years ago, when northern forests were replaced by open grassland, thus prompting an explosive evolution of small, burrowing rodents. The weasel's ancestors were larger than the current form, and underwent a reduction in size to exploit the new food source. The least weasel thrived during the Ice Age, as its small size and long body allowed it to easily operate beneath snow, as well as hunt in burrows. It probably crossed to North America through the Bering land bridge 200,000 years ago.<ref name=mac205>Template:Harvnb</ref>

SubspeciesEdit

The least weasel has a high geographic variation, a fact which has historically led to numerous disagreements among biologists studying its systematics. The least weasel's subspecies are divided into three categories:<ref name=s975>Template:Harvnb</ref>

  • The pygmaea–rixosa group (small least weasels): Tiny weasels with short tails, pedomorphic skulls, and pelts that turn pure white in winter. They inhabit northern European Russia, Siberia, the Russian Far East, Finland, the northern Scandinavian Peninsula, Mongolia, northeastern China, Japan and North America.<ref name=s975/>
  • The boccamela group (large least weasels): Very large weasels with large skulls, relatively long tails and lighter coloured pelts. Locally, they either do not turn white or only partially change colour in winter. They inhabit Transcaucasia, from western Kazakhstan to Semirechye and in the flat deserts of Middle Asia. They are also found in Morocco, Algeria, and Tunisia.<ref name=s975/>
  • The nivalis group (medium-sized least weasels): Medium-sized weasels, with tails of moderate length, representing a transitional form between the former two groups. They inhabit the middle and southern regions of European Russia, Crimea, the Ciscaucasus, western Kazakhstan, the southern and middle Urals and the montane parts of Middle Asia, except the Kopet Dag.<ref name=s975/>
Subspecies Trinomial authority Description Range Synonyms
Common least weasel
M. n. nivalis

(Nominate subspecies) File:Mustelanivalisnivalis.png

Linnaeus, 1766 A medium-sized subspecies with a tail of moderate length, constituting about 20–21% of its body length. In its summer fur, the upper body is a dark brownish or chestnut colour, while its winter fur is pure white. It is probably a transitional form between the small pygmaea and the large vulgaris.<ref name=s982>Template:Harvnb</ref> The middle regions of European Russia, from the Baltic states to the middle and southern Urals, northward approximately to the latitude of Saint Petersburg and Perm, and south to the Kursk and Voronezh Oblasts. Outside the former Soviet Union, its range includes Northern Europe (except for Ireland, Iceland, Finland and parts of the Scandinavian Peninsula) and Hokkaidō. caraftensis (Kishida, 1936)

kerulenica (Bannikov, 1952)
punctata (Domaniewski, 1926)
yesoidsuna (Kishida, 1936)

Allegheny least weasel
M. n. allegheniensis
Rhoads, 1901 Similar to M. n. rixosa, but is larger, has a broad skull and darker coat, and is more adapted to live in deciduous forests<ref>Template:Cite journal</ref> The northeastern United States (Michigan, Pennsylvania, New York, Virginia, North Carolina, Ohio, Illinois, Wisconsin, West Virginia, and Indiana)
Transcaucasian least weasel
M. n. boccamela

File:Mustela-nivalis.jpg

Bechstein, 1800 A very large subspecies, with a long tail constituting about 30% of its body length. In its summer fur, the upper body is light brownish or chestnut with yellowish or reddish tints, with some individuals having a brownish dot on the corners of the mouth and sometimes on the chest and belly. The winter fur is not pure white, being usually dirty white with brown patches.<ref name=s980>Template:Harvnb</ref> Transcaucasia, southern Europe, Asia Minor and probably western Iran italicus (Barrett-Hamilton, 1900)
Plains least weasel
M. n. campestris
Jackson, 1913 The Great Plains of the United States (South Dakota, Iowa, Nebraska, and Kansas)
Caucasian least weasel
M. n. caucasica
Barrett-Hamilton, 1900 dinniki (Satunin, 1907)
Alaskan least weasel
M. n. eskimo
Stone, 1900 A small subspecies. Resembles M. n. rixosa, but has a duller colour, a larger skull and a shorter tail.<ref>Template:Cite journal</ref> Alaska, the Yukon, and the Northwest Territories
Turkmenian least weasel
M. n. heptneri
Morozova-Turova, 1953 A very large subspecies with a long tail constituting about 25–30% of its body length. In its summer fur, the upper body is very light sandy brown or pale-yellowish. The fur is short, sparse and coarse, and does not turn white in winter.<ref name=s981>Template:Harvnb</ref> The deserts and semi-deserts of southern Kazakhstan and Middle Asia from the Caspian Sea to Semirechye, southern Tajikistan, Koppet Dag, Afghanistan and northeastern Iran
Japanese least weasel
M. n. namiyei
Kuroda, 1921 Smaller than M. n. rixosa and paler than M. n. eskimo. Resembles M. n. pygmaea, but the head and body are longer and the tail considerably longer.<ref>Template:Cite journal</ref> Northern Honshū (Aomori, Akita and Iwate Prefectures)
Mediterranean least weasel
M. n. numidica

File:Mustelanivalisiberica.png

Pucheran, 1855 The largest subspecies Morocco, Algeria, Egypt (formerly thought to be a distinct species, the Egyptian weasel), Malta, the Azores Islands and Corsica albipes (Mina Palumbo, 1868)

algiricus (Thomas, 1895)
atlas (Barrett-Hamilton, 1904)
corsicanus (Cavazza, 1908)
fulva (Mina Palumbo, 1908)
galanthias (Bate, 1905)
ibericus (Barrett-Hamilton, 1900)
meridionalis (Costa, 1869)
siculus (Barrett-Hamilton, 1900) subpalmata Hemprich & Ehrenberg, 1833

Montane Turkestan least weasel
M. n. pallida
Barrett-Hamilton, 1900 A medium-sized subspecies with a tail constituting about 24% of its body length. The colour of the summer fur is light-brownish, while the winter fur is white.<ref name="s984">Template:Harvnb</ref> The montane parts of Turkmenistan, Uzbekistan, Tajikistan, Kazakhstan and Kirgizia, as well as the Chinese parts of the same mountain systems and perhaps in the extreme eastern parts of Hindukush
Siberian least weasel
M. n. pygmaea
J. A. Allen, 1903 A very small subspecies, with a short tail which constitutes about 13% of its body length. In its summer coat, the dorsal colour is dark brown or reddish, while the winter fur is entirely white.<ref name="s978">Template:Harvnb</ref> All of Siberia (except for southern and southeastern Transbaikalia); the northern and middle Urals, northern Kazakhstan and the Russian Far East, including Sakhalin, the Kuril Islands and the Korean Peninsula; all of Mongolia (except for the eastern part), and probably northeastern China kamtschatica (Dybowksi, 1922)
Bangs' least weasel
M. n. rixosa

File:Mustela nivalis (two, fighting).jpg

Bangs, 1896 The smallest subspecies and the smallest living mammalian carnivore in the world. In its summer coat, the fur is dark reddish-brown, while the winter fur is pure white.<ref name=m14>Template:Harvnb</ref> Nunavut, Labrador, Quebec, Minnesota, North Dakota, Montana, Saskatchewan, Alberta, and British Columbia
Middle European least weasel
M. n. vulgaris
Erxleben, 1777 A somewhat larger subspecies than nivalis with a longer tail, which constitutes about 27% of its body length. In its summer fur, the upper body varies from being light brownish to dark chestnut, while the winter fur is white in its northern range and piebald in its southern range.<ref name=s983>Template:Harvnb</ref> Southern European Russia from the latitude of southern Voronezh and Kursk districts, Crimea, Ciscaucasia, and the northern slopes of the main Caucasus, eastward to the Volga. Outside the former Soviet Union, its range includes Europe southward to the Alps and the Pyrenees. Introduced to New Zealand.<ref name=long>Long, J. L. (2003). Introduced Mammals of the World: Their History, Distribution and Influence. Cabi Publishing. pp. 271–272. Template:ISBN</ref><ref>Template:Cite book</ref> dumbrowskii (Matschie, 1901)

hungarica (Vásárhelyi, 1942)
minutus (Pomel, 1853)
monticola (Cavazza, 1908)
nikolskii (Semenov, 1899)
occidentalis (Kratochvil, 1977)
trettaui (Kleinschmidt, 1937)
vasarhelyi (Kretzoi, 1942)

DescriptionEdit

File:Mustela nivalis 03 MWNH 768.jpg
Skull of a least weasel
File:Frenataermineanivalis.png
Skulls of a long-tailed weasel (top), a stoat (bottom left) and least weasel (bottom right), as illustrated in Merriam's Synopsis of the Weasels of North America

The least weasel has a thin, greatly elongated and extremely flexible body with a small, yet elongated, blunt-muzzled head which is no thicker than the neck. The eyes are small in relation to their head size and are bulging and dark colored. The legs and tail are relatively short, the latter constituting less than half the body length. The feet have sharp, dark-coloured claws, and the soles are heavily haired.<ref name=s967>Template:Harvnb</ref> The skull, especially that of the small rixosa group, has an infantile appearance when compared with that of other members of the genus Mustela (in particular, the stoat and kolonok). This is expressed in the relatively large size of the cranium and shortened facial region.<ref name=s969>Template:Harvnb</ref> The skull is, overall, similar to that of the stoat, but smaller, though the skulls of large male weasels tend to overlap in size with those of small female stoats.<ref name=h468/> There are usually four pairs of nipples but these are only visible in females.

The baculum is short,<ref>Template:Cite book</ref> Template:Cvt, with a thick, straight shaft. Fat is deposited along the spine, kidneys, gut mesentries and around the limbs. The least weasel has muscular anal glands under the tail, which measure Template:Cvt, and contain sulphurous volatiles, including thietanes and dithiacyclopentanes. The smell and chemical composition of these chemicals are distinct from those of the stoat.<ref name=h468>Template:Harvnb</ref> The least weasel moves by jumping, the distance between the tracks of the fore and hind limbs being Template:Cvt.<ref name=s991>Template:Harvnb</ref>

File:Weaselskeleton.jpg
Skeleton, as illustrated in Lydekker's The New Natural History

Dimensions vary geographically, to an extent rarely found among other mammals. Least weasels of the boccamela group, for example, may outweigh the smaller races by almost four times. In some large subspecies, the male may be 1.5 times longer than the female. The tail lengths are also variable, constituting 13 to 30 percent of the length of the body. Average body length in males is Template:Cvt, while females average Template:Cvt. The tail measures Template:Cvt in males and Template:Cvt in females. Males weigh Template:Cvt, while females weigh Template:Cvt.<ref name=s970>Template:Harvnb</ref>

File:Mustela winter.jpg
The winter coat is conspicuous when there is no snow on the ground.

The winter fur is dense, but short and closely fitting. In northern subspecies, the fur is soft and silky, but coarse in southern forms. The summer fur is very short, sparser and rougher. The upper parts in the summer fur are dark, but vary geographically from dark-tawny or dark-chocolate to light pale tawny or sandy. The lower parts, including the lower jaw and inner sides of the legs, are white. There is often a brown spot at the corner of the mouth. The dividing line between the dark upper and light lower parts is usually straight but sometimes forms an irregular line. The tail is brown, and sometimes the tip is a little darker but it is never black. In the northern part of its range and at high altitudes, the least weasel changes colour in the winter, the coat becoming pure white and exhibiting a few black hairs in rare circumstances.<ref name=s969/><ref name=Collins>Template:Cite book</ref>

Distribution and habitatEdit

File:Alaska Weasel.jpg
Alaskan least weasel (M. n. eskimo)
File:Walid soukkou 99.jpg
Least weasel in northern Algeria

The least weasel has a circumboreal, Holarctic distribution, encompassing much of Europe and North Africa, Asia and parts of northern North America. It has been introduced to New Zealand,<ref name=long/> Malta, Crete, the Azore Islands and also São Tomé off West Africa. It occurs throughout Europe, except Ireland, and on all major Mediterranean islands.<ref>Template:Cite journal</ref> It also occurs on Honshu and Hokkaido Islands in Japan and on Kunashir, Iturup, and Sakhalin Islands in Russia.<ref name=iucn/>

The least weasel inhabits fields, open woodland, bushy and rocky areas, parks and gardens at elevations of up to about Template:Cvt.<ref name=Collins/>

Fossilised remains of the least weasel are known from Denisova Cave.<ref>Template:Cite journal</ref>

Behaviour and ecologyEdit

Reproduction and developmentEdit

The least weasel mates in April–July and there is a 34- to 37-day gestation period. In the Northern Hemisphere, the average litter size consists of 6 kits and these reach sexual maturity in 3 to 4 months. Males may mate during their first year of life, though this is usually unsuccessful. They are fecund in February–October, though the early stages of spermatogenesis do occur throughout the winter months. Anestrus in females lasts from September until February.<ref name=h474>Template:Harvnb</ref>

The female raises her kits without help from the male. They are Template:Cvt in weight at birth. Newborn kits are born pink, naked, blind and deaf, but gain a white coat of downy fur at the age of 4 days. At 10 days, the margin between the dark upper parts and light under parts becomes visible. The milk teeth erupt at 2 to 3 weeks of age, at which point the young start to eat solid food, though lactation can last 12 weeks. The eyes and ears open at 3 to 4 weeks of age, and by 8 weeks, killing behaviour is developed. The family breaks up after 9 to 12 weeks.<ref name=h474/> There is a single litter each year and least weasels can live for 7 or 8 years.<ref name=Collins/>

Territorial and social behavioursEdit

File:Mustela nivalis (two, fighting).jpg
Two least weasels fighting

The least weasel has a typical mustelid territorial pattern, consisting of exclusive male ranges encompassing multiple female ranges. The population density of each territory depends greatly on food supply and reproductive success, thus the social structure and population density of any given territory is unstable and flexible.<ref>Template:Cite journal</ref> Like the stoat, the male least weasel extends his range during spring or during food shortages. Its scent marking behaviour is similar to that of the stoat; it uses faeces, urine and anal and dermal gland secretions, the latter two of which are deposited by anal dragging and body rubbing. The least weasel does not dig its own den, but nests in the abandoned burrow of another species such as a mole or rat.<ref name=h471>Template:Harvnb</ref> The burrow entrance measures about Template:Cvt across and leads to the nest chamber located up to Template:Cvt below ground. The nest chamber (which is used for sleeping, rearing kits and storing food) measures Template:Cvt in diameter, and is lined with straw and the skins of the weasel's prey.<ref name=m277>Template:Harvnb</ref>

The least weasel has four basic vocalisations; a guttural hiss emitted when alarmed, which is interspersed with short screaming barks and shrieks when provoked. When defensive, it emits a shrill wail or squeal. During encounters between males and females or between a mother and kits, the least weasel emits a high-pitched trilling. The least weasel's way of expressing aggression is similar to that of the stoat. Dominant weasels exhibit lunges and shrieks during aggressive encounters, while subdominant weasels will emit submissive squeals.<ref name=h471/>

DietEdit

File:Donnola vs lepre.JPG
Taxidermy exhibit showing a least weasel attacking a European hare, in the Natural History Museum of Genoa

The least weasel feeds predominantly on mouse-like rodents, including mice, hamsters, gerbils and others. It usually does not attack adult hamsters and rats. Frogs, fish, small birds and bird eggs are rarely eaten. It can deal with adult pikas and gerbils, but usually cannot overcome brown rats and sousliks. Exceptional cases are known of least weasels killing prey far larger than themselves, such as capercaillie, hazel hen and hares.<ref name=s987>Template:Harvnb</ref> In England, a favoured prey item is the field vole (Microtus agrestis). These have fluctuations in population size, and in years of abundance may form up to 54% of the weasel's diet. In years of scarcity, birds form a greater proportion of the diet and female least weasels may fail to breed.<ref>Template:Cite journal</ref>

Despite its small size, the least weasel is a fierce hunter, capable of killing a rabbit five to 10 times its own weight.<ref name=mac208>Template:Harvnb</ref> Although they are commonly taken, the rabbits are usually young specimens, and become an important food source during the spring, when small rodents are scarce and rabbit kits are plentiful. Male least weasels take a higher proportion of rabbits than females, as well as an overall greater variety of prey. This is linked to the fact that being larger, and having vaster territorial ranges than females, males have more opportunities to hunt a greater diversity of prey.<ref name=h472>Template:Harvnb</ref>

The least weasel forages undercover, to avoid being seen by foxes and birds of prey. It is adapted for pursuing its prey down tunnels, though it may also bolt prey from a burrow and kill it in the open.<ref name=h472/> The least weasel kills small prey, such as voles, with a bite to the occipital region of the skull<ref name=s987/> or the neck, dislocating the cervical vertebrae. Large prey typically dies of blood loss or circulatory shock.<ref name=h472/> When food is abundant, only a small portion of the prey is eaten, usually the brain. The average daily food intake is Template:Cvt, which is equivalent to 30–35% of the animal's body weight.<ref name=s987/>

Predators and competitorsEdit

File:Weaselsvsstoat.png
Least weasels driven from a mountain hare carcass by a stoat, as illustrated in Barrett-Hamilton's A History of British Mammals

The least weasel is small enough to be preyed upon by a range of other predators.<ref name=h475>Template:Harvnb</ref> Least weasel remains have been found in the excrement of red foxes, sables, steppe and forest polecat, stoats, eagle owls and buzzards.<ref name=s992>Template:Harvnb</ref> The owls most efficient at capturing least weasels are barn, barred, and great horned owls. Other birds of prey threatening to the least weasel include broad-winged and rough-legged buzzards. Some snake species may prey on the least weasel, including the black rat snake and copperhead.<ref name=m277/> Aside from its smaller size, the least weasel is more vulnerable to predation than the stoat because it lacks a black predator deflection mark on the tail.<ref name=h475/>

In areas where the least weasel is sympatric with the stoat, the two species compete with each other for rodent prey. The weasel manages to avoid too much competition by living in more upland areas, feeding on smaller prey and being capable of entering smaller holes. It actively avoids encounters with stoats, though female weasels are less likely to stop foraging in the presence of stoats, perhaps because their smaller size allows them to quickly escape into holes.<ref name=h469>Template:Harvnb</ref>

Diseases and parasitesEdit

Ectoparasites known to infest weasels include the louse Trichodectes mustelae and the mites Demodex and Psoregates mustela. The species may catch fleas from the nests and burrows of its prey. Flea species known to infest weasels include Ctenophthalmus bisoctodentatus and Palaeopsylla m. minor, which they get from moles, P. s. soricis, which they get from shrews, Nosopsyllus fasciatus, which they get from rodents and Dasypsyllus gallinulae which they get from birds.<ref name=h475/>

Helminths known to infest weasels include the trematode Alaria, the nematodes Capillaria, Filaroides and Trichinella and the cestode Taenia.<ref name=h475/> Least weasels are commonly infected with the nematode Skrjabingylus nasicola, adults of which are found in the nasal sinuses and can damage the skull. There is no evidence that this has serious detrimental effects on even heavily infested animals.<ref>Template:Cite journal</ref>

ConservationEdit

The least weasel is listed as Least Concern on the IUCN Red List, because of its wide global distribution and large population that is thought to be not in decline.<ref name=iucn/>

In folklore and mythologyEdit

Macedonian and Greek cultureEdit

File:Wenceslas Hollar - The basilisk and the weasel.jpg
17th century print of a least weasel confronting a basilisk

The Ancient Macedonians believed that to see a least weasel was a good omen. In some districts of Macedon, women who suffered from headaches after having washed their heads in water drawn overnight would assume that a weasel had previously used the water as a mirror, but they would refrain from mentioning the animal's name for fear that it would destroy their clothes.<ref name="Abbott">Abbott, G. A. (1903), Macedonian Folklore, pp. 108–109, Cambridge University Press</ref>

Similarly, a popular superstition in southern Greece had it that the least weasel had previously been a bride, who was transformed into a bitter animal which would destroy the wedding dresses of other brides out of jealousy.<ref name="Abbott"></ref> According to Pliny the Elder, the least weasel was the only animal that was capable of killing the basilisk:

To this dreadful monster the effluvium of the weasel is fatal, a thing that has been tried with success, for kings have often desired to see its body when killed; so true is it that it has pleased Nature that there should be nothing without its antidote. The animal is thrown into the hole of the basilisk, which is easily known from the soil around it being infected. The weasel destroys the basilisk by its odour, but dies itself in this struggle of nature against its own self.<ref>{{#invoke:citation/CS1|citation

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Ojibwe and Inuit cultureEdit

The Ojibwe believed that the least weasel could kill the dreaded wendigo by rushing up its anus.<ref>Barnouw, Victor (1979). Wisconsin Chippewa Myths & Tales: And Their Relation to Chippewa Life, p. 53, University of Wisconsin Press, Template:ISBN</ref>

In Inuit mythology, the least weasel is credited with both great wisdom and courage, and whenever a mythical Inuit hero wished to accomplish a valorous task, he would generally change himself into a least weasel.<ref>Dufresne, Frank (2005), Alaska's Animals and Fishes, p. 109, Kessinger Publishing, Template:ISBN</ref>

See alsoEdit

ReferencesEdit

CitationsEdit

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General and cited referencesEdit

Further readingEdit

External linksEdit

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