View source for Biology:Stoat

{{short description|Species of mammal (mustelid)}}
{{speciesbox
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name=iucn>{{cite iucn |title=''Mustela erminea'' |author=Reid, F. |author2=Helgen, K. |author3=Kranz, A. |date=2016 |page=e.T29674A45203335 |doi=10.2305/IUCN.UK.2016-1.RLTS.T29674A45203335.en |access-date=19 November 2021}}</ref>
| image = Mustela erminea upright.jpg
| genus = Mustela
| species = erminea
| authority = [[Biography:Carl Linnaeus|Linnaeus]], [[Biology:10th edition of Systema Naturae|1758]]
| range_map = Stoat area.png
| range_map_caption = Stoat range (includes ''M. richardsonii'' and ''M. haidarum'')
{{leftlegend|green|native|outline=gray}}
{{leftlegend|red|introduced|outline=gray}}
}}

The '''stoat''' ('''''Mustela erminea'''''), also known as the '''Eurasian ermine''', '''Beringian ermine''' and '''ermine''', is a mustelid native to [[Earth:Eurasia|Eurasia]] and the northern regions of [[Earth:North America|North America]]. Because of its wide [[Biology:Circumpolar distribution|circumpolar distribution]], it is listed as Least Concern on the [[Organization:IUCN Red List|IUCN Red List]].<ref name=iucn/> It was [[Biology:Stoat in New Zealand|introduced into New Zealand]] in the late 19th century to control rabbits, but had a devastating effect on native bird populations and was nominated as one of the world's top 100 "worst invaders".<ref>{{cite web |title=100 of the World's Worst Invasive Species |url = http://www.issg.org/database/species/ecology.asp?si=98&fr=1&sts=&lang=EN |publisher=Invasive Species Specialist Group}}</ref>

The name ermine ({{IPAc-en|ˈ|ɜːr|m|ɪ|n}}) is used for species in the genus ''[[Biology:Mustela|Mustela]]'', especially the stoat, in its pure white winter coat, or the fur thereof.<ref name=SOED>{{cite book|title=Shorter Oxford English dictionary|year= 2007|publisher=Oxford University Press |location= UK |isbn= 978-0199206872|pages=3804}}</ref>
Ermine fur was used in the 15th century by Catholic monarchs, who sometimes used it as the mozzetta cape. It has long been used on the ceremonial robes of members of the United Kingdom House of Lords. It was also used in capes on images such as the [[Religion:Infant Jesus of Prague|Infant Jesus of Prague]].

==Etymology==
[[File:Mustela erminea 02 MWNH 382e.jpg|thumb|Skull]]
The root word for "stoat" is likely either the Dutch word {{lang|nl|stout}} ("bold")<ref name="c124"/> or the [[Social:Gothic language|Gothic]] word {{lang|got|𐍃𐍄𐌰𐌿𐍄𐌰𐌽}} ({{lang|got-Latn|stautan}}, "to push").<ref name="j160">{{Harvnb|Johnston|1903|p=160}}</ref> According to John Guillim, in his ''Display of Heraldrie'', the word "ermine" is likely derived from Armenia, the nation where it was thought the species originated,<ref name="c124">{{Harvnb|Coues|1877|pp=124–125}}</ref> though other authors have linked it to the Norman French from the Teutonic {{lang|gem|harmin}} (Anglo-Saxon {{lang|ang|hearma}}). This seems to come from the [[Social:Lithuanian language|Lithuanian]] word {{lang|lt|šarmu}}.<ref name="j160"/> In Ireland (where the [[Biology:Least weasel|least weasel]] does not occur), the stoat is referred to as a weasel, while in North America it is called a short-tailed weasel. A male stoat is called a ''dog'', ''hob'', or ''jack'', while a female is called a ''jill''. The collective noun for stoats is either ''gang'' or ''pack''.<ref name="h456">{{Harvnb|Harris|Yalden|2008|p=456}}</ref>

== Taxonomy ==
Formerly considered a single species with a very wide circumpolar range, a 2021 study split ''M. erminea'' into three species: ''M. erminea sensu stricto'' (Eurasia and northern North America), ''M. richardsonii'' (most of North America), and ''M. haidarum'' (several islands off the [[Place:Pacific Northwest|Pacific Northwest]] coast).<ref name=Colella_al2021>{{Cite journal|last1=Colella|first1=Jocelyn P. |last2=Frederick |first2=Lindsey M. |last3=Talbot|first3=Sandra L.|last4=Cook|first4=Joseph A. |date=2021|title=Extrinsically reinforced hybrid speciation within Holarctic ermine (''Mustela'' spp.) produces an insular endemic |journal=Diversity and Distributions |volume=27|issue=4|pages=747–762 |doi=10.1111/ddi.13234 |s2cid=234059194 |doi-access=}}</ref><ref>{{Cite journal |last1=Fleming |first1=Melissa A.|last2=Cook|first2=Joseph A. |date=2002|title=Phylogeography of endemic ermine (''Mustela erminea'') in southeast Alaska |doi=10.1046/j.1365-294X.2002.01472.x |journal=Molecular Ecology |volume=11|issue=4|pages=795–807 |pmid=11972765|s2cid=18874091}}</ref><ref name=":1">{{Cite web|title=Explore the Database|url=https://www.mammaldiversity.org/explore.html#species-id=1005863|access-date=2021-07-12|website=www.mammaldiversity.org}}</ref>

===Subspecies===
{{As of|2021}}, 21 [[Biology:Subspecies|subspecies]] are recognized.<ref name=Colella_al2021/>
{{Clear}}
{| class="wikitable" style="width:100%;"
|- style="background:#115a6c;"
!Subspecies
!Trinomial authority
!Description
!Range
!Synonyms
|-
|Northern stoat ''M. e. erminea''
(Nominate subspecies)
|Linnaeus, 1758
|A small-to-medium-sized subspecies with a relatively short and broad facial region<ref name="s1010">{{Harvnb|Heptner|Sludskii|2002|p=1010}}</ref>
|The Kola Peninsula, [[Place:Scandinavia|Scandinavia]]
|<small>''hyberna'' (Kerr, 1792)</small><br />
<small>''maculata'' (Billberg, 1827)</small><br />
|-
|Middle Russian stoat<br />''M. e. aestiva''
[[File:Mustela.erminea.jpg|150 px]]
|Kerr, 1792
|A moderately sized subspecies with dark, tawny or chestnut summer fur<ref name="s1010"/>
|European Russia (except for the Kola Peninsula), [[Place:Central Europe|Central]] and [[Place:Western Europe|Western Europe]]
|<small>''algiricus'' (Thomas, 1895)</small><br />
<small>''alpestris'' (Burg, 1920)</small><br />
<small>''giganteus'' (Burg, 1920)</small><br />
<small>''major'' (Nilsson, 1820)</small>
|-
|Tundra stoat ''M. e. arctica''
[[File:Mustelaerminaarcticaskull.png|150 px]]
|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<ref name="m15">{{Harvnb|Merriam|1896|p=15}}</ref>
|[[Earth:Alaska|Alaska]], northwestern Canada, and the [[Place:Arctic Archipelago|Arctic Archipelago]] (except for Baffin Island)
|<small>''audax'' (Barrett-Hamilton, 1904)</small><br />
<small>''kadiacensis'' (Merriam, 1896)</small><br />
<small>''kadiacensis'' (Osgood, 1901)</small><br />
<small>''richardsonii'' (Bonaparte, 1838)</small><br />
|-
|''M. e. augustidens''
|Brown, 1908
|
|
|
|-
|Fergana stoat ''M. e. ferghanae''
|Thomas, 1895
|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.<ref name="s1014">{{Harvnb|Heptner|Sludskii|2002|p=1014}}</ref><ref>{{Cite journal |author1=Kotia, A. |author2=Angmo, K. |author3=Bharti, R. R. |author4=Adhikari, B. S.  |author5=Rawat, G. S. |date=2011 |title=A record of the little-known Stoat ''Mustela erminea ferghanae'' from Ladakh, India |journal=Small Carnivore Conservation |volume=44 |pages=42–43 |url=https://www.researchgate.net/figure/Himalayan-race-of-Stoat-Mustela-erminea-ferghanae-with-prey-10-July-2010-Zanskar_fig1_270339150}}</ref>
|Tien Shan and Pamir-Alay mountains, Afghanistan, India, western Tibet and the adjacent parts of the Tien Shan in China
|<small>''shnitnikovi'' (Ognev, 1935)</small><br/>
<small>''whiteheadi'' (Wroughton, 1908)</small>
|-
|Irish stoat ''M. e. hibernica''
[[File:Mustela erminea hibernica.jpg|150 px]]
|Thomas and Barrett-Hamilton, 1895
|Larger than ''aestiva'', but smaller than ''stabilis''. 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.<ref name="h459">{{Harvnb|Harris|Yalden|2008|p=459}}</ref>
|Ireland and the Isle of Man
|
|-
|Kodiak stoat ''M. e. kadiacensis''
|Merriam, 1896
|
|Kodiak Island
|
|-
|East Siberian stoat (known locally as Ezo stoat in Japan)<br />''M. e. kaneii''
|Baird, 1857
|A moderately sized subspecies. It is smaller than ''M. e. tobolica'', with close similarities to ''M. e. arctica''. The colour of the summer coat is relatively light, with varying intensities of browning-yellow tinges.<ref name="s1012"/>
|Eastern Siberia and the [[Place:Russian Far East|Russian Far East]] including Kamchatka, except the Amur Oblast and Ussuriland, Transbaikalia and the Sayan Mountains. Also in Hokkaidō.
|<small>''baturini'' (Ognev, 1929)</small><br/>
<small>''digna'' (Hall, 1944)</small><br/>
<small>''kamtschatica'' (Dybowski, 1922)</small><br/>
<small>''kanei'' (G. Allen, 1914)</small><br/>
<small>''naumovi'' (Jurgenson, 1938)</small><br/>
<small>''orientalis'' (Ognev, 1928)</small><br/>
<small>''transbaikalica'' (Ognev, 1928)</small>
|-
|Karaginsky stoat ''M. e. karaginensis''
|Jurgenson, 1936
|A very small subspecies with a light chestnut-coloured summer coat<ref name="s1013">{{Harvnb|Heptner|Sludskii|2002|p=1013}}</ref>
|Karaginsky Island, along the eastern coast of Kamchatka
|
|-
|Altai stoat ''Mustela e. lymani''
|Hollister, 1912
|A moderately sized subspecies with less dense fur than ''M. e. tobolica''. The colour of its summer coat consists of weakly developed reddish-brown tones. The skull is similar to that of ''M. e. aestiva''.<ref name="s1012">{{Harvnb|Heptner|Sludskii|2002|p=1012}}</ref>
|The mountains of southern Siberia eastwards to [[Earth:Lake Baikal|Baikal]] and the contiguous parts of Mongolia
|
|-
|''M. e. martinoi''
|Ellerman and Morrison-Scott, 1951
|
|
|<small>''birulai'' (Martino and Martino, 1930)</small><br />
|-
|Swiss stoat<br />''M. e. minima''
|Cavazza, 1912
|
|Switzerland
|
|-
|Gobi stoat
''M. e. mongolica''
|Ognev, 1928
|
|The Govi-Altai Province
|
|-
|Japanese stoat ''M. e. nippon''
[[File:Mustela erminea in summer.jpg|150 px]]
|Cabrera, 1913
|
|northern Honshū
|
|-
|
''M. e. ognevi''
|Jurgenson, 1932
|
|
|
|-
|Polar stoat ''M. e. polaris''
|Barrett-Hamilton, 1904
|
|[[Place:Greenland|Greenland]]
|
|-
|Hebrides stoat ''M. e. ricinae''
|Miller, 1907
|
|The Hebrides
|
|-
|''M. e. salva''
|Hall, 1944
|
|
|
|-
|British stoat ''M. e. stabilis''
[[File:Mustela Erminea head.jpg|150 px]]
|Barrett-Hamilton, 1904
|Larger than mainland European stoats<ref name="h459"/>
|Great Britain; introduced to New Zealand
|
|-
|Caucasian stoat ''M. e. teberdina''
|Korneev, 1941
|A small subspecies with a coffee to reddish-tawny summer coat<ref name="s1010"/>
|The northern slope of the middle part of the main [[Earth:Caucasus|Caucasus]] range
|<small>''balkarica'' (Basiev, 1962)</small>
|-
|Tobolsk stoat ''M. e. tobolica''
|Ognev, 1923
|A large subspecies; it is somewhat larger than ''aestiva'', with long and dense fur.<ref name="s1011">{{Harvnb|Heptner|Sludskii|2002|p=1011}}</ref>
|Western Siberia, eastwards to the Yenisei and Altai Mountains and in Kazakhstan
|
|}

==Evolution==
The stoat's direct ancestor was ''Mustela palerminea'', a common carnivore in central and eastern Europe during the [[Earth:Middle Pleistocene|Middle Pleistocene]],<ref name="k101">{{Harvnb|Kurtén|1968|pp=101–102}}</ref> that spread to North America during the late [[Biology:Blancan|Blancan]] or early [[Biology:Irvingtonian|Irvingtonian]].<ref name="k150">{{Harvnb|Kurtén|1980|p=150}}</ref> 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 explosive [[Biology:Evolution|evolution]] 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 the [[Biology:Long-tailed weasel|long-tailed weasel]], which is in a different genus (''[[Biology:Neogale|Neogale]]''), arose as its mirror image in North America 2 million years ago. The stoat thrived during the [[Earth:Last Glacial Period|Ice 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 falling [[Sea level|sea level]]s exposed the Bering land bridge.<ref name="mac205">{{Harvnb|Macdonald|1992|p=205}}</ref>

Combined [[Biology:Phylogenetics|phylogenetic]] analyses indicate the stoat's closest living relatives are the American ermine (''M. richardsonii'') and Haida ermine (''M. haidarum''), the latter of which partially descends from ''M. erminea''.<ref name=Colella_al2021/> It is basal to most other members of ''Mustela'', with only the yellow-bellied (''M. kathia''), Malayan (''M. katiah''), and back-striped (''M. strigidorsa'') weasels being more basal.<ref name="Law-2018">{{Cite journal|last1=Law|first1=C. J.|last2=Slater|first2=G. J.|last3=Mehta|first3=R. S.|date=2018-01-01|title=Lineage Diversity and Size Disparity in Musteloidea: Testing Patterns of Adaptive Radiation Using Molecular and Fossil-Based Methods|journal=Systematic Biology|volume=67|issue=1|pages=127–144|doi=10.1093/sysbio/syx047|pmid=28472434|doi-access=free}}</ref> The mountain 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 genus ''[[Biology:Neogale|Neogale]]'' such as the long-tailed weasel, but as those species have since been separated into a new genus, this is likely not the case.<ref name="h458">{{Harvnb|Harris|Yalden|2008|p=458}}</ref>

==Description==
===Build===
[[File:Stoat and weasel (cropped).jpg|thumb|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{{clarify|date=March 2023|reason=define what this means: including tail?}},<ref name="s997">{{Harvnb|Heptner|Sludskii|2002|p=997}}</ref> 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 little
more than half its length.<ref name="c117">{{Harvnb|Coues|1877|pp=117–121}}</ref> The skull, although very similar to that of the least weasel, is relatively longer, with a narrower braincase. The projections of the skull and teeth are weakly developed, but stronger than those of the least weasel.<ref name="s999">{{Harvnb|Heptner|Sludskii|2002|p=999}}</ref> The eyes are round, black and protrude slightly. The [[Biology:Whiskers|whiskers]] 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 curved [[Biology:Baculum|baculum]] with a proximal knob that increases in weight as it ages.<ref name="h457"/> [[Chemistry:Fat|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.<ref name="h457">{{Harvnb|Harris|Yalden|2008|p=457}}</ref>

[[File:Weasel (Mustela erminea).jpg|thumb|Skeleton]]
The dimensions of the stoat are variable, but not as significantly as the least weasel's.<ref name="s1002"/> Unusual among the Carnivora, the size of stoats tends to decrease proportionally with latitude, in contradiction to [[Physics:Bergmann's rule|Bergmann's rule]].<ref name="k101"/> [[Biology:Sexual dimorphism|Sexual dimorphism]] in size is pronounced, with males being roughly 25% larger than females and 1.5-2.0 times their weight.<ref name="h459"/> On average, males measure {{cvt|187|–|325|mm}} in body length, while females measure {{cvt|170|–|270|mm}}. The tail measures {{cvt|75|–|120|mm}} in males and {{cvt|65|–|106|mm}} in females. In males, the hind foot measures {{cvt|40.0|-|48.2|mm}}, while in females it is {{cvt|37.0|-|47.6|mm}}. The height of the ear measures {{cvt|18.0|-|23.2|mm}} in males and {{cvt|14.0|-|23.3|mm}}. The skulls of males measure {{cvt|39.3|-|52.2|mm}} in length, while those of females measure {{cvt|35.7|-|45.8|mm}}. Males average {{cvt|258|g}} in weight, while females weigh less than {{cvt|180|g}}.<ref name="s1002">{{Harvnb|Heptner|Sludskii|2002|p=1002}}</ref>

The stoat has large anal [[Biology:Scent gland|scent gland]]s measuring {{cvt|8.5|x|5|mm}} in males and smaller in females. Scent glands are also present on the cheeks, belly and flanks.<ref name="h457"/> [[Biology:Epidermis (zoology)|Epidermal]] secretions, which are deposited during body 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 its [[Biology:Anal gland|anal gland]]s, giving rise to a strong, musky odour produced by several sulphuric compounds. The odour is distinct from that of least weasels.<ref name="h460"/>

===Fur===
[[File:Mustela erminea winter cropped.jpg|thumb|A stoat in winter fur]]
The winter fur is very dense and silky, but quite closely lying and short, while the summer fur is rougher, shorter and sparse.<ref name="s997"/> 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 by photoperiod, 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.<ref name="h457"/> Differences in the winter and summer coats are less apparent in southern forms of the species.<ref name="s998">{{Harvnb|Heptner|Sludskii|2002|p=998}}</ref> In the species' southern range, the coat remains brown, but is denser and sometimes paler than in summer.<ref name="h457"/>

==Distribution and habitat==
The stoat has a circumboreal 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, [[Place:Svalbard|Svalbard]], the Mediterranean islands and some small North Atlantic islands. In Japan, it is present in central mountains (northern and central Japan Alps) to northern part of Honshu (primarily above 1,200 m) and Hokkaido. Its vertical range is from sea level to {{cvt|3000|m}}.<ref name=iucn/> In North America, it is found throughout [[Earth:Alaska|Alaska]] and western [[Earth:Yukon|Yukon]] to most of Arctic Canada east to [[Place:Greenland|Greenland]]. Throughout the rest of North America, as well as parts of [[Earth:Nunavut|Nunavut]] including Baffin Island and some islands in southeast Alaska, it is replaced by ''M. richardsonii''.<ref name=Colella_al2021/>

===Introduction to New Zealand===
{{main|Biology:Stoats in New Zealand}}
Stoats were introduced into [[Earth:New Zealand|New 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 ornithologist Walter 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.<ref>{{cite book |last=King |first=Carolyn |title=Immigrant Killers |publisher=Oxford University Press |location=Auckland, NZ |year=1984 |isbn=0-19-558121-0 }}</ref> 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 southern beechmast (beechnuts), which enable the reproduction of rodents on which stoats also feed, enabling stoats to increase their own numbers.<ref>{{cite journal |last1=Purdey |first1=D. C. |first2=C. M. |last2=King |first3=B. |last3=Lawrence |year=2004 |title=Age structure, dispersion and diet of a population of stoats (''Mustela erminea'') in southern Fiordland during the decline phase of the beechmast cycle |journal=New Zealand Journal of Zoology |publisher=The Royal Society of New Zealand |volume=31 |pages=205–225 |url=http://researchcommons.waikato.ac.nz/bitstream/10289/230/1/content.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://researchcommons.waikato.ac.nz/bitstream/10289/230/1/content.pdf |archive-date=2022-10-09 |url-status=live |access-date=2009-11-30 |doi=10.1080/03014223.2004.9518373 |issue=3 |s2cid=55061896 |doi-access=free }}</ref> For instance, the endangered [[Biology:South Island takahē|South Island takahē]]'s wild population dropped by a third between 2006 and 2007, after a stoat plague triggered by the 2005–06 [[Biology:Mast (botany)|mast]] wiped out more than half the takahē in untrapped areas.<ref>{{cite news |url=http://www.stuff.co.nz/environment/298858 |title=Stoats decimating takahe in Fiordland |date=4 March 2008 |work=stuff.co.nz |access-date=23 April 2011 }}</ref>

==Behaviour and ecology==
===Reproduction and development===
[[File:Mustela.erminea.young.jpg|thumb|Young stoat]]
In the Northern Hemisphere, mating occurs in the April–July period. In spring, the male's testes are enlarged, a process accompanied by an increase of [[Chemistry:Testosterone|testosterone]] concentration in the [[Biology:Blood plasma|plasma]]. [[Biology:Spermatogenesis|Spermatogenesis]] occurs in December, and the males are fertile from May to August, after which the testes regress.<ref>Gulamhusein, A. P., and W. H. Tam. "[https://rep.bioscientifica.com/downloadpdf/journals/rep/41/2/jrf_41_2_006.pdf Reproduction in the male stoat, Mustela erminea]." Reproduction 41.2 (1974): 303-312.</ref> Female stoats are usually only in heat for a brief period, which is triggered by changes in day length.<ref name="k215">{{Harvnb|King|Powell|2007|pp=215}}</ref> [[Biology:Copulation (zoology)|Copulation]] can last as long as 1 hour.<ref>Amstislavsky, Sergei, and Yulia Ternovskaya. "[https://web.archive.org/web/20171117174933/https://pdfs.semanticscholar.org/f3b8/69a6701e7bb11035119faf36745f0bd0fafc.pdf Reproduction in mustelids]." Animal Reproduction Science 60 (2000): 571-581.</ref> Stoats are not [[Biology:Monogamy in animals|monogamous]], with litters often being of mixed paternity. Stoats undergo [[Biology:Embryonic diapause|embryonic 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.<ref name="k209">{{Harvnb|King|Powell|2007|pp=209–210}}</ref> The gestation 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.<ref name="k215"/> Females can reabsorb embryos and in the event of a severe winter they may reabsorb their entire litter.<ref name="k255">{{Harvnb|King|Powell|2007|pp=255}}</ref> Males play no part in rearing the young, which are born blind, deaf, toothless and covered in fine white or pinkish down. The milk 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. [[Biology:Lactation|Lactation]] ends after 12 weeks. Prior to the age of five to seven weeks, kits have poor [[Physics:Thermoregulation|thermoregulation]], 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 being weaned.<ref name="h464">{{Harvnb|Harris|Yalden|2008|pp=464–465}}</ref>

===Territorial and sheltering behaviour===
[[File:StoatDen3 (cropped).jpg|thumb|Stoat nesting in a hollow tree.]]
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 sexes mark their territories with [[Biology:Urine|urine]], faeces and two types of scent marks; anal drags are meant to convey territorial occupancy, and body rubbing is associated with agonistic encounters.<ref name="h460">{{Harvnb|Harris|Yalden|2008|pp=460–461}}</ref>

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 in [[Biology:Magpie|magpie]] nests. Males and females typically live apart, but close to each other.<ref name="s1021">{{Harvnb|Heptner|Sludskii|2002|pp=1021–1022}}</ref> Each stoat has several dens dispersed within its range. A single den has several galleries, mainly within {{cvt|30|cm}} of the surface.<ref name="h461">{{Harvnb|Harris|Yalden|2008|p=461}}</ref>

===Diet===
As with the least weasel, mouse-like rodents predominate in the stoat's diet. It regularly preys on larger rodent and lagomorph species, and takes individuals far larger than itself. In Russia, its prey includes rodents and lagomorphs such as [[Biology:European water vole|European water vole]]s, common hamsters, pikas and others, which it overpowers in their burrows. Prey species of secondary importance include small [[Biology:Bird|bird]]s, [[Biology:Fish|fish]], and shrews and, more rarely, [[Biology:Amphibian|amphibian]]s, [[Biology:Lizard|lizard]]s, and [[Biology:Insect|insect]]s.<ref name="s1018">{{Harvnb|Heptner|Sludskii|2002|p=1018}}</ref> It also preys on [[Biology:Lemming|lemming]]s.<ref>{{cite news |title=It's Feast Or Famine: Predators May Drive Lemming Cycles, Science Researchers Say |date=2003 |author=American Association For The Advancement Of Science |publisher=ScienceDaily |url=https://www.sciencedaily.com/releases/2003/11/031104063521.htm#:~:text=The%20results%20showed%20the%20stoats,exclusively%20on%20lemmings%20for%20prey}}</ref>
In Great Britain, [[Biology:European rabbit|European rabbit]]s 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 the [[Biology:Myxomatosis|myxomatosis]] 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, [[Biology:Rat|rat]]s, 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. [[Biology:Hare|Hare]]s are sometimes taken, but are usually young specimens.<ref name="h463"/> In New Zealand, the stoat feeds principally on birds, including the rare [[Biology:Kiwi (bird)|kiwi]], kaka, [[Biology:Yellowhead (bird)|mohua]], [[Biology:Yellow-crowned parakeet|yellow-crowned parakeet]], and [[Biology:New Zealand dotterel|New Zealand dotterel]].<ref name="h463">{{Harvnb|Harris|Yalden|2008|p=463}}</ref> Cases are known of stoats preying on young [[Biology:Muskrat|muskrat]]s. The stoat typically eats about {{cvt|50|g}} of food a day, which is equivalent to 25% of the animal's live weight.<ref name="s1020">{{Harvnb|Heptner|Sludskii|2002|p=1020}}</ref>

[[File:Stoat killing a rabbit.jpg|thumb|Stoat killing a European rabbit]]
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 the weasel war dance), though this behaviour could be linked to ''[[Biology:Metastrongylidae|Skrjabingylus]]'' infections.<ref name="h463"/> The stoat seeks to immobilize large prey such as rabbits with a bite to the spine at the back of the neck. The stoat may [[Biology:Surplus killing|surplus kill]] when the opportunity arises, though excess prey is usually cached and eaten later to avoid obesity, as overweight stoats tend to be at a disadvantage when pursuing prey into their burrows.<ref name="v417">{{Harvnb|Verts|Carraway|1998|p=417}}</ref> Small prey typically die instantly from a bite to the back of the neck, while larger prey, such as rabbits, typically die of [[Medicine:Shock (circulatory)|shock]], as the stoat's canine teeth are too short to reach the spinal column or major arteries.<ref name="h463"/>

===Communication===
The stoat is a usually silent animal, but 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.<ref name="h460"/>

Aggressive behavior in stoats is categorized in these forms:<ref name="h460"/>
* 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
* [[Biology:Kleptoparasitism|Kleptoparasitism]], in which a dominant stoat appropriates the killing 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.<ref name="h460"/>

===Predators===
Larger mammalian predators such as [[Biology:Red fox|red fox]]es (''Vulpes vulpes'') and sables (''Martes zibellina'') are known to prey on stoats.<ref name="s1025">{{Harvnb|Heptner|Sludskii|2002|p=1025}}</ref> Additionally, a wide range of birds of prey can take stoats, from small [[Biology:Northern hawk-owl|northern hawk-owl]]s (''Surnia ulula'') and [[Biology:Short-eared owl|short-eared owl]]s (''Asio flammeus'') to various [[Biology:Buteo|buzzards]], [[Biology:Milvus|kites]], [[Biology:Goshawk|goshawk]]s, and  even Eurasian eagle-owls (''Bubo bubo'') and golden eagles (''Aquila chrysaetos'').<ref>Korpimäki, Erkki, and Kai Norrdahl. "Avian predation on mustelids in Europe 1: occurrence and effects on body size variation and life traits." Oikos (1989): 205-215.</ref> Although not classified as birds of prey, grey herons (''Ardea cinerea'') are known to prey on stoats.<ref>SAWARA, Yuji, Muneki SAKUYAMA, and Gen DEMACHI. "Diets and foraging site utilization of the Grey Heron, Ardea cinerea, in the breeding season." Japanese Journal of Ornithology 43.2 (1994): 61-71.</ref>

===Diseases and parasites===
[[Medicine:Tuberculosis|Tuberculosis]] has been recorded in stoats inhabiting the former Soviet Union and New Zealand. They are largely resistant to [[Medicine:Tularemia|tularemia]], but are reputed to suffer from [[Biology:Canine distemper|canine distemper]] in captivity. Symptoms of mange have also been recorded.<ref name="h466">{{Harvnb|Harris|Yalden|2008|p=466}}</ref>

Stoats are vulnerable to ectoparasites associated with their prey and the nests of other animals on which they do not prey. The [[Medicine:Louse|louse]] ''Trichodectes erminea'' is recorded in stoats living in Canada, Ireland and New Zealand. In continental Europe, 26 flea species are recorded to infest stoats, including ''Rhadinospylla pentacantha'', ''Megabothris rectangulatus'', ''Orchopeas howardi'', ''Spilopsyllus ciniculus'', ''Ctenophthalamus nobilis'', ''[[Biology:Moorhen flea|Dasypsyllus gallinulae]]'', ''Nosopsyllus fasciatus'', ''Leptospylla segnis'', ''Ceratophyllus gallinae'', ''Parapsyllus n. nestoris'', ''Amphipsylla kuznetzovi'' and ''Ctenopsyllus bidentatus''. [[Biology:Tick|Tick]] species known to infest stoats are ''Ixodes canisuga'', ''[[Biology:Ixodes hexagonus|I. hexagonus]]'', and ''[[Biology:Ixodes ricinus|I. ricinus]]'' and ''[[Biology:Haemaphysalis longicornis|Haemaphysalis longicornis]]''. Louse species known to infest stoats include ''Mysidea picae'' and ''Polyplax spinulosa''. [[Biology:Mite|Mite]] species known to infest stoats include ''[[Biology:Neotrombicula|Neotrombicula autumnalis]]'', ''[[Biology:Demodex|Demodex erminae]]'', ''[[Biology:Eulaelaps|Eulaelaps stabulans]]'', ''Gymnolaelaps annectans'', ''Hypoaspis nidicorva'', and ''[[Biology:Listrophorus|Listrophorus mustelae]]''.<ref name="h466"/>

The [[Biology:Nematode|nematode]] ''[[Biology:Skrjabingylus nasicola|Skrjabingylus nasicola]]'' is particularly threatening to stoats, as it erodes the bones of the nasal sinuses and decreases fertility. Other nematode species known to infect stoats include ''Capillaria putorii'', ''Molineus patens'' and ''Strongyloides martes''. Cestode species known to infect stoats include ''[[Biology:Taenia (tapeworm)|Taenia tenuicollis]]'', ''Mesocestoides lineatus'' and rarely ''[[Biology:Acanthocephala|Acanthocephala]]''.<ref name="h466"/>

==In culture==
[[File:Lady with an Ermine - Leonardo da Vinci (adjusted levels).jpg|thumb|Leonardo da Vinci's ''Lady with an Ermine'' (1489–1491).<br> National Museum, Kraków, Poland.]]

===Folklore and mythology===
In Irish mythology, stoats were viewed anthropomorphically 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.<ref>Monaghan, Patricia (2004) ''The encyclopedia of Celtic mythology and folklore: Facts on File library of religion and mythology'', page 426, Infobase Publishing, {{ISBN|0-8160-4524-0}}</ref> Stoats were also supposed to hold the souls of infants who died before [[Religion:Baptism|baptism]].<ref>Daniels, Cora Linn & Stevans, C. M. ''Encyclopedia of Superstitions, Folklore, and the Occult Sciences of the World, Volume 2'' (2003), The Minerva Group, Inc., {{ISBN|1-4102-0915-6}}</ref> 

In the folklore of the Komi people of the Urals, stoats are symbolic of beautiful and coveted young women.<ref>Laakso, Johanna (2005) ''Our otherness: Finno-Ugrian approaches to women's studies, or vice versa, Volume 2 of Finno-Ugrian studies in Austria'', LIT Verlag Münster, {{ISBN|3-8258-8626-3}}</ref> In the [[Religion:Zoroastrianism|Zoroastrian]] 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.<ref>Sax, Boria (2001) ''The mythical zoo: an encyclopedia of animals in world myth, legend, and literature'', ABC-CLIO, {{ISBN|1-57607-612-1}}</ref> 

The former nation (now region) of Brittany in France uses a stylized ermine-fur pattern in forming the coat of arms and flag of Brittany. Gilles Servat's song ''La Blanche Hermine'' ("The White Ermine") became an anthem for Bretons (and is popular among French people in general).

In the 16th century Chinese novel ''Investiture of the Gods'', [[Unsolved:Erlang Shen|Erlang Shen]] transforms into an ermine to demonstrate his shapeshifting abilities.{{cn |date=March 2023}}

===Fur use===

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 the traditional ancient symbol of the Duchy of Brittany, forming its earliest flag. There is also a design called ermine inspired by the winter coat of the stoat and painted onto other furs, such as rabbit.<ref name="lords">{{cite news | url = http://news.bbc.co.uk/2/hi/special_report/1999/01/99/lords_reform/235653.stm | title = A house of traditions | date = January 19, 1999 | work=BBC News}}</ref> In Europe these furs are a symbol of royalty and high status. The ceremonial robes of members of the United Kingdom House of Lords and the academic hoods of the universities of Oxford and Cambridge are traditionally trimmed with ermine.<ref name="lords"/> In practice, rabbit or fake fur is now often used due to expense or [[Philosophy:Animal rights|animal 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'', by [[Biography:Leonardo da Vinci|Leonardo da Vinci]]. Henry Peacham's ''Emblem 75'', which depicts an ermine being pursued by a hunter and two hounds, is entitled "Cui 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.<ref>[http://f01.middlebury.edu/FS010A/STUDENTS/n075.htm The Minerva Britanna Project] {{Webarchive|url=https://web.archive.org/web/20030418124136/http://f01.middlebury.edu/FS010A/students/n075.htm |date=2003-04-18 }}</ref>

Ermine (both ''M. erminea'' and ''M. richardsonii'', both of which inhabited the Tlingit's territory) were also valued by the Tlingit and other indigenous 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.<ref>{{cite web|url=http://www.corbisimages.com/stock-photo/rights-managed/IH011277/tlingit-ermineskin-shirt-daa-dugu-koodas|title=Tlingit Ermine-Skin Shirt (Daa dugu k'oodas')}}</ref>

The stoat was a fundamental item in the fur trade of the [[Place:Soviet Union|Soviet Union]], with no less than half the global catch coming from within its borders. The Soviet Union also contained the highest grades of stoat pelts, with the best grade North American pelts being comparable only to the 9th grade in the quality criteria of former Soviet stoat standards. Stoat harvesting never became a specialty in any Soviet republic, with most stoats being captured incidentally in traps or near villages. Stoats in the Soviet Union were captured either with dogs or with box-traps or jaw-traps. Guns were rarely used, as they could damage the pelt.<ref name="s1029">{{Harvnb|Heptner|Sludskii|2002|pp=1029–1030}}</ref>

<gallery widths="160px" heights="200px">
File:Marvel Rea (left), Ford Sterling, and Alice Maison, appearing in Mack Sennett Comedies LCCN89711780.jpg|American actress Alice Maison shown wearing ermine fur in a Mack Sennett comedy film
File:Thea Sternheim c. 1910, by Franz Grainer.jpg|Thea Sternheim, wife of playwright Carl Sternheim, wearing an ermine hat
</gallery>

==References==
=== Citations ===
{{Reflist|colwidth=30em}}

===Bibliography===
{{refbegin|40em}}
* {{Cite book|last=Ahern|first=Albert|url=https://archive.org/details/furfacts00aherrich|title=Fur Facts|location=St. Louis|publisher=C. P. Curran printing company|year=1922}}
* {{Cite book|last=Coues|first=Elliott|url=https://archive.org/details/furbearinganima00couegoog|title=Fur-bearing Animals: A Monograph of North American Mustelidae|publisher=Government Printing Office|year=1877}}
* {{Cite book|last1=Harris|first1=Stephen|last2=Yalden|first2=Derek|title=Mammals of the British Isles|publisher=Mammal Society|edition=4th Revised|year=2008|isbn=978-0-906282-65-6}}
* {{Cite book|last=Johnston|first=Harry Hamilton|url=https://archive.org/details/britishmammalsat00john|title=British mammals; an attempt to describe and illustrate the mammalian fauna of the British islands from the commencement of the Pleistocene period down to the present day|publisher=London, Hutchinson|year=1903}}
* {{cite book |last1=King |first1=Carolyn M. |first2=Roger A.|last2=Powell |title=The Natural History of Weasels and Stoats: Ecology, Behavior, and Management |url=https://books.google.com/books?id=In48DwAAQBAJ |edition=2nd |year=2007 |others=Illustrations by Consie Powell |location=Oxford, UK |publisher=Oxford University Press |isbn=978-0-19-530056-7 |oclc=228781387}}
* {{Cite book|last=Kurtén|first=Björn|title=Pleistocene mammals of Europe|publisher=Weidenfeld and Nicolson|year=1968}}
* {{Cite book|last=Kurtén|first=Björn|title=Pleistocene mammals of North America|publisher=Columbia University Press|year=1980|isbn=0-231-03733-3}}
* {{Cite book|last1=Heptner|first1=V. G.|last2=Sludskii|first2=A. A.|url=https://archive.org/details/mammalsofsov212001gept|title=Mammals of the Soviet Union. Vol. II, part 1b, Carnivores (Mustelidae and Procyonidae) |location=Washington, D.C. |publisher=Smithsonian Institution Libraries and National Science Foundation|year=2002|isbn=90-04-08876-8}}
* {{Cite book|last=Macdonald|first=David|url=https://archive.org/details/velvetclawnatura00macd|title=The Velvet Claw: A Natural History of the Carnivores|location=New York|publisher=Parkwest|year=1992|isbn=0-563-20844-9|url-access=registration}}
* {{Cite book|last=Merriam|first=Clinton Hart|url=https://archive.org/details/synopsisofweasel00merriala|title=Synopsis of the weasels of North America |location=Washington, D.C. |publisher=Govt. Print. Off.|year=1896}}
* {{Cite book|last1=Verts|first1=B. J.|last2=Carraway|first2=Leslie N.|url=https://books.google.com/books?id=8KI1AmzIDnwC&pg=PA415 |title=Land Mammals of Oregon|publisher=University of California Press|year=1998|isbn=0-520-21199-5}}
{{refend}}

==External links==
* [https://web.archive.org/web/20081013032427/http://www.arkive.org/species/ARK/mammals/Mustela_erminea/ Erminea (''Mustela erminea'')] at [[ARKive]]
* [http://www.funet.fi/pub/sci/bio/life/mammalia/carnivora/mustelidae/mustela/index.html#erminea ''Mustela erminea'' taxonomy]
* [https://web.archive.org/web/20140617054359/http://www.doc.govt.nz/conservation/threats-and-impacts/animal-pests/animal-pests-a-z/stoats/you-can-help/ Stoat control information]
* [http://news.bbc.co.uk/1/hi/uk/7875271.stm Stoat 'playing'(?) in snow]
* [https://web.archive.org/web/20100603192205/http://www.doc.govt.nz/publications/conservation/land-and-freshwater/offshore-islands/secretary-island-restoration-project/ Fiordland Islands NZ stoat eradication]
* [http://www.bbc.co.uk/nature/species/Stoat BBC Wildlife finder including video footage and sound files]
* [http://www.stevenround-birdphotography.com/Stoat.htm Stoat images] {{Webarchive|url=https://web.archive.org/web/20120731152703/http://www.stevenround-birdphotography.com/Stoat.htm |date=2012-07-31 }}
* [http://www.mnh.si.edu/mna/image_info.cfm?species_id=191 Smithsonian Institution—North American Mammals: ''Mustela erminea'']
* [http://siwild.si.edu/content/animal-groups/other-carnivores/ermine.htm?species=149 Smithsonian Wild: ''Mustela erminea'']

{{Taxonbar|from=Q25345}}

[[Category:Arctic land animals]]
[[Category:Carnivorans of Asia]]
[[Category:Carnivorans of Europe]]
[[Category:Carnivorans of North America]]

{{Sourceattribution|Stoat}}