Editing Wolf

{{Short description|Species of canine}}
{{Redirect2|Grey Wolf|Gray Wolf|other uses|Grey Wolf (disambiguation)|and|Wolf (disambiguation)}}
{{featured article}}
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{{Use mdy dates|date=December 2019}}
{{Use Canadian English|date=December 2019}}
{{Speciesbox
| name = Wolf
| fossil_range = {{longitem|style=line-height:1.25em|{{nowrap|[[Middle Pleistocene]] – present}} {{nowrap|(400,000–0 [[Before Present|YBP]])}}}}
| image = Eurasian wolf 2.jpg
| image_caption = {{longitem|[[Eurasian wolf]] (''Canis lupus lupus'') at [[Polar Park (animal park)|Polar Park]] in Bardu, Norway}}
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 2 June 2024">{{cite iucn |author=Boitani, L. |author2=Phillips, M. |author3=Jhala, Y. |name-list-style=amp |year=2023 |title=''Canis lupus'' |amends=2018 |page=e.T3746A247624660 |doi=10.2305/IUCN.UK.2023-1.RLTS.T3746A247624660.en |access-date=2 June 2024}}</ref>
| status2 = CITES_A2
| status2_system = CITES
| status2_ref = <ref>{{Cite web|title=Appendices {{!}} CITES|url=https://cites.org/eng/app/appendices.php|access-date=2022-01-14|website=cites.org}}</ref>{{efn|Populations of Bhutan, India, Nepal, and Pakistan are included in Appendix I. Excludes domesticated form and dingo, which are referenced as ''Canus lupus familiaris'' and ''Canus lupus dingo''.}}
| taxon = Canis lupus
| authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]<ref name=Linnaeus1758/>
| subdivision_ranks = Subspecies
| subdivision = {{small|See [[subspecies of Canis lupus|Subspecies of ''Canis lupus'']]}}
| range_map = Canis lupus distribution (IUCN).png
| range_map_caption = Global wolf range based on IUCN's 2023 assessment.<ref name="iucn status 2 June 2024" />
}}

The '''wolf''' ('''''Canis lupus''''';{{efn|Domestic and feral [[dog]]s are included in the [[phylogenetic]] but not colloquial definition of 'wolf', and are thus not in the scope of this article.}} {{plural form}}: '''wolves'''),<!--This article uses "wolf" rather than "grey wolf" or "gray wolf" throughout; see the discussion at https://en.wikipedia.org/wiki/Talk:Wolf/Archive_6#Requested_move_2_August_2018--> also known as the '''grey wolf''' or '''gray wolf''', is a [[Canis|canine]] native to [[Eurasia]] and [[North America]]. More than thirty [[subspecies of Canis lupus|subspecies of ''Canis lupus'']] have been recognized, including the [[dog]] and [[dingo]], though grey wolves, as popularly understood, only comprise [[Wild type|naturally-occurring wild]] subspecies. The wolf is the largest wild [[Neontology|extant]] member of the family [[Canidae]], and is further distinguished from other ''[[Canis]]'' species by its less pointed ears and muzzle, as well as a shorter torso and a longer tail. The wolf is nonetheless related closely enough to smaller ''Canis'' species, such as the [[coyote]] and the [[golden jackal]], to produce fertile [[Canid hybrid|hybrids]] with them. The wolf's fur is usually mottled white, brown, grey, and black, although subspecies in the arctic region may be nearly all white.

Of all members of the [[genus]] ''Canis'', the wolf is most [[Generalist and specialist species|specialized]] for [[Pack hunter|cooperative]] [[Hunting behavior of gray wolves|game hunting]] as demonstrated by its physical adaptations to tackling large prey, its more [[Sociality|social nature]], and its highly advanced [[Wolf communication|expressive behaviour]], including individual or group [[howling]]. It travels in [[Nuclear family|nuclear families]] consisting of a [[mated pair]] accompanied by their offspring. Offspring may leave to form their own [[Pack (canine)|pack]]s on the onset of sexual maturity and in response to competition for food within the pack. Wolves are also [[Territory (animal)|territorial]], and fights over territory are among the principal causes of mortality. The wolf is mainly a [[carnivore]] and feeds on large wild [[hooved mammal]]s as well as smaller animals, livestock, [[carrion]], and garbage. Single wolves or mated pairs typically have higher success rates in hunting than do large packs. [[Pathogen]]s and parasites, notably the [[rabies|rabies virus]], may infect wolves.

The global wild wolf population was estimated to be 300,000 in 2003 and is considered to be of [[Least-concern species|Least Concern]] by the [[International Union for Conservation of Nature]] (IUCN). Wolves have a long history of interactions with humans, having been despised and hunted in most [[Pastoralism|pastoral]] communities because of their attacks on livestock, while conversely being respected in some [[Agrarian society|agrarian]] and [[hunter-gatherer]] societies. Although the fear of wolves exists in many human societies, the majority of recorded attacks on people have been attributed to animals suffering from [[rabies]]. [[Wolf attack]]s on humans are rare because wolves are relatively few, live away from people, and have developed a fear of humans because of their experiences with hunters, farmers, ranchers, and shepherds.

== Etymology ==

{{See also|Wolf (name)}}

The English "wolf" stems from the [[Old English]] {{wikt-lang|ang|wulf}}, which is itself thought to be derived from the [[Proto-Germanic language|Proto-Germanic]] {{lang|gem-x-proto|*wulfaz}}. The [[Proto-Indo-European root]] *''{{PIE|[[wikt:Reconstruction:Proto-Indo-European/wĺ̥kʷos|wĺ̥kʷos]]}}'' may also be the source of the [[Latin]] word for the animal ''lupus'' (*''{{PIE|lúkʷos}}'').<ref>{{OEtymD|wolf}}</ref><ref name=Lehrman/> The name "grey wolf" refers to the greyish colour of the species.<ref name=Goldman/>

Since pre-Christian times, [[Germanic peoples]] such as the [[Anglo-Saxon]]s took on ''wulf'' as a [[prefix]] or [[suffix]] in their names. Examples include Wulfhere ("Wolf Army"), Cynewulf ("Royal Wolf"), Cēnwulf ("Bold Wolf"), Wulfheard ("Wolf-hard"), Earnwulf ("Eagle Wolf"), Wulfstān ("Wolf Stone") Æðelwulf ("Noble Wolf"), Wolfhroc ("Wolf-Frock"), Wolfhetan ("Wolf Hide"), Scrutolf ("Garb Wolf"), Wolfgang ("Wolf Gait") and Wolfdregil ("Wolf Runner").{{sfn|Marvin|2012|pp=74–75}}

== Taxonomy ==
{{cladogram|title=Canine phylogeny with ages of divergence
|caption=Cladogram and divergence of the grey wolf (including the domestic dog) among its closest extant relatives<ref name=Koepfli-2015/>
|cladogram={{clade| style=font-size:85%;line-height:75%;width:475px;
|sublabel1=''3.50 [[Million years ago|mya]]''<!--E-->
|1={{clade
   |sublabel1=''3.06 mya''<!--F-->
   |1={{clade
      |sublabel1=''2.74 mya''<!--G-->
      |1={{clade
         |sublabel1=''1.92 mya''<!--H-->
         |1={{clade
            |sublabel1=''1.62 mya''<!--I-->
            |1={{clade
               |sublabel1=''1.32 mya''<!--J-->
               |1={{clade
                  |sublabel1=''1.10 mya''<!--K-->
                  |1={{clade
                     |1='''Grey wolf''' [[File:Dogs, jackals, wolves, and foxes (Plate I).png|50 px]]
                     |2=[[Coyote]] [[File:Dogs, jackals, wolves, and foxes (Plate IX).png|50 px]]
                     }}
                  |2=[[African wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate XI).jpg|50 px]]
                  }}
               |2=[[Ethiopian wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate VI).png|50 px]]
               }}
            |2=[[Golden jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate X).png|50 px]]
            }}
         |2=[[Dhole]] [[File:Dogs, jackals, wolves, and foxes (Plate XLI).png|50 px]]
         }}
      |2=[[African wild dog]] [[File:Dogs, jackals, wolves, and foxes (Plate XLIV).png|50 px]]
      }}
   |sublabel2=''2.62 mya''<!--D-->
   |2={{clade
      |1=[[Side-striped jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate XIII).png|50 px]]
      |2=[[Black-backed jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate XII).png|50 px]]
      }}
   }}
}}
}}
In 1758, the Swedish botanist and zoologist [[Carl Linnaeus]] published in his ''[[10th edition of Systema Naturae|Systema Naturae]]'' the [[binomial nomenclature]].<ref name=Linnaeus1758/> ''[[Canis]]'' is the Latin word meaning "[[dog]]",<ref>{{OEtymD|canine}}</ref> and under this [[genus]] he listed the doglike carnivores including domestic dogs, wolves, and [[jackal]]s. He classified the domestic dog as ''Canis familiaris'', and the wolf as ''Canis lupus''.<ref name=Linnaeus1758/> Linnaeus considered the dog to be a separate species from the wolf because of its "cauda recurvata" (upturning tail) which is not found in any other [[canid]].<ref name=Clutton-Brock1995/>

=== Subspecies ===
{{Main|Subspecies of Canis lupus}}
{{Further|Pleistocene wolf}}
In the third edition of ''[[Mammal Species of the World]]'' published in 2005, the [[mammalogist]] [[W. Chris Wozencraft|W. Christopher Wozencraft]] listed under ''C. lupus'' 36 wild subspecies, and proposed two additional subspecies: ''familiaris'' (Linnaeus, 1758) and ''[[Canis lupus dingo|dingo]]'' (Meyer, 1793). Wozencraft included ''hallstromi''—the [[New Guinea singing dog]]—as a [[taxonomic synonym]] for the [[dingo]]. Wozencraft referred to a 1999 [[mitochondrial DNA]] (mtDNA) study as one of the guides in forming his decision, and listed the 38 [[Subspecies of Canis lupus|subspecies of ''C. lupus'']] under the biological [[common name]] of "wolf", the [[nominate subspecies]] being the [[Eurasian wolf]] (''C. l. lupus'') based on the [[type specimen]] that Linnaeus studied in Sweden.<ref name=Wozencraft2005/> Studies using [[Paleogenomics|paleogenomic]] techniques reveal that the modern wolf and the dog are [[sister taxa]], as modern wolves are not closely related to the population of wolves that was first [[Domestication|domesticated]].<ref name=Larson2014/> In 2019, a workshop hosted by the [[IUCN]]/Species Survival Commission's Canid Specialist Group considered the New Guinea singing dog and the dingo to be [[feral dog|feral]] ''Canis familiaris'', and therefore should not be assessed for the [[IUCN Red List]].<ref name=Alvares2019/>

=== Evolution ===
{{Main|Evolution of the wolf}}
{{Further|Domestication of the dog}}
[[File:Canis mosbachensis Wikipedia Juandertal.jpg|thumb|right|Life restoration of ''[[Canis mosbachensis]]'', the wolf's immediate ancestor]]
The [[phylogenetic]] descent of the extant wolf ''C. lupus'' from the earlier ''[[Canis mosbachensis|C. mosbachensis]]'' (which in turn descended from ''[[Canis etruscus|C. etruscus]]'') is widely accepted.{{sfn|Mech|Boitani|2003|pp=239–245}} Among the oldest fossils of the modern grey wolf is from Ponte Galeria in Italy, dating to 406,500 ± 2,400 years ago.<ref name=":2">{{Cite journal |last1=Iurino |first1=Dawid A. |last2=Mecozzi |first2=Beniamino |last3=Iannucci |first3=Alessio |last4=Moscarella |first4=Alfio |last5=Strani |first5=Flavia |last6=Bona |first6=Fabio |last7=Gaeta |first7=Mario |last8=Sardella |first8=Raffaele |date=2022-02-25 |title=A Middle Pleistocene wolf from central Italy provides insights on the first occurrence of Canis lupus in Europe |journal=Scientific Reports |language=en |volume=12 |issue=1 |page=2882 |doi=10.1038/s41598-022-06812-5 |issn=2045-2322 |pmc=8881584 |pmid=35217686|bibcode=2022NatSR..12.2882I }}</ref> Remains from Cripple Creek Sump in Alaska may be considerably older, around 1 million years old,<ref name=Tedford2009/> though differentiating between the remains of modern wolves and ''C. mosbachensis'' is difficult and ambiguous, with some authors choosing to include C. ''mosbachensis'' (which first appeared around 1.4 million years ago) as an early subspecies of ''C. lupus.''<ref name=":2" />

Considerable morphological diversity existed among wolves by the [[Late Pleistocene]]. Many Late Pleistocene wolf populations had more robust skulls and teeth than modern wolves, often with a shortened [[Rostrum (anatomy)#Vertebrates|snout]], a pronounced development of the [[temporalis]] muscle, and robust [[premolar]]s. It is proposed that these features were specialized adaptations for the processing of carcass and bone associated with the hunting and scavenging of [[Pleistocene megafauna]]. Compared with modern wolves, some Pleistocene wolves showed an increase in tooth breakage similar to that seen in the extinct [[dire wolf]]. This suggests they either often processed carcasses, or that they competed with other carnivores and needed to consume their prey quickly. The frequency and location of tooth fractures in these wolves indicates they were habitual bone crackers like the modern [[spotted hyena]].<ref name=Thalmann2018/>

[[Genome|Genomic]] studies suggest modern wolves and dogs descend from a common ancestral wolf population.<ref name=Freedman2014/><ref name=Skoglund2015/><ref name=Fan2016/> A 2021 study found that the [[Himalayan wolf]] and the [[Indian wolf|Indian plains wolf]] are part of a [[Lineage (evolution)|lineage]] that is [[Basal (phylogenetics)|basal]] to other wolves and [[Cladogenesis|split]] from them 200,000 years ago.<ref name=Hennelly2021/> Other wolves appear to share most of their common ancestry much more recently, within the last 23,000 years (around the peak and the end of the [[Last Glacial Maximum]]), originating from [[Siberia]]<ref name=":0">{{Cite journal |last1=Bergström |first1=Anders |last2=Stanton |first2=David W. G. |last3=Taron |first3=Ulrike H. |last4=Frantz |first4=Laurent |last5=Sinding |first5=Mikkel-Holger S. |last6=Ersmark |first6=Erik |last7=Pfrengle |first7=Saskia |last8=Cassatt-Johnstone |first8=Molly |last9=Lebrasseur |first9=Ophélie |last10=Girdland-Flink |first10=Linus |last11=Fernandes |first11=Daniel M. |last12=Ollivier |first12=Morgane |last13=Speidel |first13=Leo |last14=Gopalakrishnan |first14=Shyam |last15=Westbury |first15=Michael V. |date=2022-07-14 |title=Grey wolf genomic history reveals a dual ancestry of dogs |journal=Nature |language=en |volume=607 |issue=7918 |pages=313–320 |doi=10.1038/s41586-022-04824-9 |issn=0028-0836 |pmc=9279150 |pmid=35768506|bibcode=2022Natur.607..313B }}</ref> or [[Beringia]].<ref name=":1">{{Cite journal |last1=Loog |first1=Liisa |last2=Thalmann |first2=Olaf |last3=Sinding |first3=Mikkel-Holger S. |last4=Schuenemann |first4=Verena J. |last5=Perri |first5=Angela |last6=Germonpré |first6=Mietje |last7=Bocherens |first7=Herve |last8=Witt |first8=Kelsey E. |last9=Samaniego Castruita |first9=Jose A. |last10=Velasco |first10=Marcela S. |last11=Lundstrøm |first11=Inge K. C. |last12=Wales |first12=Nathan |last13=Sonet |first13=Gontran |last14=Frantz |first14=Laurent |last15=Schroeder |first15=Hannes |date=May 2020 |title=Ancient DNA suggests modern wolves trace their origin to a Late Pleistocene expansion from Beringia |journal=Molecular Ecology |language=en |volume=29 |issue=9 |pages=1596–1610 |doi=10.1111/mec.15329 |issn=0962-1083 |pmc=7317801 |pmid=31840921|bibcode=2020MolEc..29.1596L }}</ref> While some sources have suggested that this was a consequence of a [[population bottleneck]],<ref name=":1" /> other studies have suggested that this a result of [[gene flow]] homogenising ancestry.<ref name=":0" />

A 2016 genomic study suggests that Old World and New World wolves split around 12,500 years ago followed by the [[Genetic divergence|divergence]] of the lineage that led to dogs from other Old World wolves around 11,100–12,300 years ago.<ref name=Fan2016/> An extinct [[Megafaunal wolf|Late Pleistocene wolf]] may have been the ancestor of the dog,<ref name=Freedman2017/><ref name=Thalmann2018/> with the dog's similarity to the extant wolf being the result of [[genetic admixture]] between the two.<ref name=Thalmann2018/> The dingo, [[Basenji]], [[Tibetan Mastiff]] and Chinese indigenous breeds are basal members of the domestic dog clade. The divergence time for wolves in Europe, the Middle East, and Asia is estimated to be fairly recent at around 1,600 years ago. Among New World wolves, the [[Mexican wolf]] diverged around 5,400 years ago.<ref name=Fan2016/>

=== Admixture with other canids ===

{{Main|Canid hybrid}}
[[File:F1 wolf-dog hybrids from Wildlife Park Kadzidlowo, Poland.png|thumb|upright=1.5|alt=Photographs of two wolf–dog hybrids standing outdoors on snowy ground|[[Wolfdog|Wolf–dog hybrids]] in the wild animal park at [[Kadzidłowo]], Poland. Left: product of a male wolf and a female [[spaniel]]; right: from a female wolf and a male [[West Siberian Laika]]]]

In the distant past, there was [[gene flow]] between [[African wolf|African wolves]], [[golden jackal]]s, and grey wolves. The African wolf is a descendant of a genetically admixed canid of 72% wolf and 28% Ethiopian wolf ancestry. One African wolf from the Egyptian [[Sinai Peninsula]] showed admixture with Middle Eastern grey wolves and dogs.<ref name=Gopalakrishnan2018/> There is evidence of gene flow between golden jackals and Middle Eastern wolves, less so with European and Asian wolves, and least with North American wolves. This indicates the golden jackal ancestry found in North American wolves may have occurred before the divergence of the Eurasian and North American wolves.<ref name=Sinding2018/>

The common ancestor of the coyote and the wolf is admixed with a [[ghost population]] of an extinct unidentified canid. This canid was genetically close to the [[dhole]] and evolved after the divergence of the [[African hunting dog]] from the other canid species. The basal position of the [[coyote]] compared to the wolf has been proposed to be due to the coyote retaining more of the mitochondrial genome of this unidentified canid.<ref name=Gopalakrishnan2018/> Similarly, a museum specimen of a wolf from southern China collected in 1963 showed a genome that was 12–14% admixed from this unknown canid.<ref name=Wang2019/> In North America, some coyotes and wolves show varying degrees of past [[Coywolf|genetic admixture]].<ref name=Sinding2018/>

In more recent times, some male [[Italian wolf|Italian wolves]] originated from dog ancestry, which indicates female wolves will breed with male dogs in the wild.<ref name=Iacolina2010/> In the [[Caucasus Mountains]], ten percent of dogs including [[livestock guardian dog]]s, are first generation hybrids.<ref name=Kopaliani2014/> Although mating between golden jackals and wolves has never been observed, evidence of [[jackal-wolf hybrid]]ization was discovered through mitochondrial DNA analysis of jackals living in the Caucasus Mountains<ref name=Kopaliani2014/> and in Bulgaria.<ref name=Moura2013/> In 2021, a genetic study found that the dog's similarity to the extant grey wolf was the result of substantial dog-into-wolf [[gene flow]], with little evidence of the reverse.<ref name="Bergström2020"/>

== Description ==

[[File:Front view of a resting Canis lupus ssp.jpg|thumb|upright|alt=Photograph of a reclining North American wolf looking directly at the photographer|A North American wolf]]

The wolf is the largest extant member of the family [[Canidae]],<ref name=Mech1974/> and is further distinguished from coyotes and jackals by a broader snout, shorter ears, a shorter torso and a longer tail.{{sfn|Heptner|Naumov|1998|pp=129–132}}<ref name=Mech1974/> It is slender and powerfully built, with a large, deeply descending [[rib cage]], a sloping back, and a heavily muscled neck.{{sfn|Heptner|Naumov|1998|p=166}} The wolf's legs are moderately longer than those of other canids, which enables the animal to move swiftly, and to overcome the deep snow that covers most of its geographical range in winter,{{sfn|Mech|1981|p=13}} though more short-legged ecomorphs are found in some wolf populations.<ref>{{Cite journal |last1=Tomiya |first1=Susumu |last2=Meachen |first2=Julie A. |date=17 January 2018 |title=Postcranial diversity and recent ecomorphic impoverishment of North American gray wolves |journal=[[Biology Letters]] |language=en |volume=14 |issue=1 |pages=20170613 |doi=10.1098/rsbl.2017.0613 |issn=1744-9561 |pmc=5803591 |pmid=29343558 }}</ref> The ears are relatively small and triangular.{{sfn|Heptner|Naumov|1998|p=166}} The wolf's head is large and heavy, with a wide forehead, strong jaws and a long, blunt muzzle.{{sfn|Heptner|Naumov|1998|pp=164–270}} The skull is {{cvt|230|–|280|mm}} in length and {{cvt|130|–|150|mm}} in width.{{sfn|Mech|1981|p=14}} The teeth are heavy and large, making them better suited to crushing bone than those of other canids, though they are not as specialized as those found in [[hyena]]s.<ref name=Therrien2005/>{{sfn|Mech|Boitani|2003|p=112}} Its [[Molar (tooth)|molars]] have a flat chewing surface, but not to the same extent as the coyote, whose diet contains more vegetable matter.<ref name=Paquet2003/> Females tend to have narrower muzzles and foreheads, thinner necks, slightly shorter legs, and less massive shoulders than males.{{sfn|Lopez|1978|p=23}}
[[File:Canis lupus italicus skeleton (white background).jpg|thumb|left|alt=Photograph of a wolf skeleton|A wolf skeleton housed in the Wolf Museum, [[Abruzzo National Park]], Italy]]

Adult wolves measure {{cvt|105|–|160|cm}} in length and {{cvt|80|–|85|cm}} at shoulder height.{{sfn|Heptner|Naumov|1998|pp=164–270}} The tail measures {{cvt|29|–|50|cm}} in length, the ears {{cvt|90|-|110|mm}} in height, and the hind feet are {{cvt|220|-|250|mm}}.{{sfn|Heptner|Naumov|1998|p=174}} The size and weight of the modern wolf increases proportionally with latitude in accordance with [[Bergmann's rule]].<ref name=Miklosi2015/> The mean body mass of the wolf is {{cvt|40|kg}}, the smallest specimen recorded at {{cvt|12|kg}} and the largest at {{cvt|79.4|kg}}.<ref name=Macdonald2001/>{{sfn|Heptner|Naumov|1998|pp=164–270}} On average, European wolves weigh {{cvt|38.5|kg}}, North American wolves {{cvt|36|kg}}, and Indian and [[Arabian wolf|Arabian wolves]] {{cvt|25|kg}}.{{sfn|Lopez|1978|p=19}} Females in any given wolf population typically weigh {{cvt|5|–|10|lb|kg}} less than males. Wolves weighing over {{cvt|54|kg}} are uncommon, though exceptionally large individuals have been recorded in Alaska and Canada.{{sfn|Lopez|1978|p=18}} In central Russia, exceptionally large males can reach a weight of {{cvt|69|-|79|kg}}.{{sfn|Heptner|Naumov|1998|p=174}}
{{Clear}}

=== Pelage ===

[[File:Tibetan Wolf Canis lupus chanko cropped.jpg|alt=Picture of a wolf standing on snowy terrain, turning its head at the camera|thumb|Wolf in [[Spiti Valley]], northern India]]

The wolf has very dense and fluffy winter fur, with a short [[Down hair|undercoat]] and long, coarse [[guard hair]]s.{{sfn|Heptner|Naumov|1998|pp=164–270}} Most of the undercoat and some guard hairs are shed in spring and grow back in autumn.{{sfn|Lopez|1978|p=19}} The longest hairs occur on the back, particularly on the front quarters and neck. Especially long hairs grow on the shoulders and almost form a crest on the upper part of the neck. The hairs on the cheeks are elongated and form tufts. The ears are covered in short hairs and project from the fur. Short, elastic and closely adjacent hairs are present on the limbs from the [[elbow]]s down to the [[calcaneal tendon]]s.{{sfn|Heptner|Naumov|1998|pp=164–270}} The winter fur is highly resistant to the cold. Wolves in northern climates can rest comfortably in open areas at {{convert|-40|C|F|abbr=on}} by placing their muzzles between the rear legs and covering their faces with their tail. Wolf fur provides better insulation than dog fur and does not collect ice when warm breath is condensed against it.{{sfn|Lopez|1978|p=19}}

In cold climates, the wolf can reduce the flow of blood near its skin to conserve body heat. The warmth of the foot pads is regulated independently from the rest of the body and is maintained at just above [[Frostbite|tissue-freezing]] point where the pads come in contact with ice and snow.{{sfn|Lopez|1978|pp=19–20}} In warm climates, the fur is coarser and scarcer than in northern wolves.{{sfn|Heptner|Naumov|1998|pp=164–270}} Female wolves tend to have smoother furred limbs than males and generally develop the smoothest overall coats as they age. Older wolves generally have more white hairs on the tip of the tail, along the nose, and on the forehead. Winter fur is retained longest by lactating females, although with some hair loss around their teats.{{sfn|Lopez|1978|p=23}} Hair length on the middle of the back is {{cvt|60|–|70|mm}}, and the guard hairs on the shoulders generally do not exceed {{cvt|90|mm}}, but can reach {{cvt|110|–|130|mm}}.{{sfn|Heptner|Naumov|1998|pp=164–270}}
[[File:Black and White Wolves.jpg|thumb|left|alt=Photograph showing one black and one white wolf standing alongside each other|Wolves in the [[La Boissière-du-Doré]] Zoo, France]]

A wolf's coat colour is determined by its guard hairs. Wolves usually have some hairs that are white, brown, grey and black.<ref name=Gipson2002/> The coat of the Eurasian wolf is a mixture of [[ochre]]ous (yellow to orange) and [[Rust (color)|rusty]] ochreous (orange/red/brown) colours with light grey. The muzzle is pale ochreous grey, and the area of the lips, cheeks, chin, and throat is white. The top of the head, forehead, under and between the eyes, and between the eyes and ears is grey with a reddish film. The neck is ochreous. Long, black tips on the hairs along the back form a broad stripe, with black hair tips on the shoulders, upper chest and rear of the body. The sides of the body, tail, and outer limbs are a pale dirty ochreous colour, while the inner sides of the limbs, belly, and groin are white. Apart from those wolves which are pure white or black, these tones vary little across geographical areas, although the patterns of these colours vary between individuals.{{sfn|Heptner|Naumov|1998|pp=168–169}}

In North America, the coat colours of wolves follow [[Gloger's rule]], wolves in the Canadian arctic being white and those in southern Canada, the U.S., and Mexico being predominantly grey. In some areas of the [[Rocky Mountains]] of Alberta and British Columbia, the coat colour is predominantly black, some being blue-grey and some with silver and black.<ref name=Gipson2002/> Differences in coat colour between sexes is absent in Eurasia;{{sfn|Heptner|Naumov|1998|p=168}} females tend to have redder tones in North America.{{sfn|Lopez|1978|p=22}} [[Black wolf|Black-coloured wolves]] in North America acquired their colour from wolf-dog admixture after the first arrival of dogs across the Bering Strait 12,000 to 14,000 years ago.<ref name=Anderson2009/> Research into the inheritance of white colour from dogs into wolves has yet to be undertaken.<ref name=Hedrick2009/>

== Ecology ==
=== Distribution and habitat ===

{{Main|Wolf distribution}}
[[File:Lupo in Sassoferrato.jpg|thumb|alt=Photograph of a wolf standing on snowy ground|An [[Italian wolf]] in a mountainous habitat in the [[Apennine Mountains|Apennines]] in [[Sassoferrato]], Italy]]

Wolves occur across Eurasia and North America. However, deliberate human persecution because of livestock predation and fear of attacks on humans has reduced the wolf's range to about one-third of its historic range; the wolf is now [[extirpated]] (locally extinct) from much of its range in Western Europe, the United States and Mexico, and completely in the [[British Isles]] and Japan. In modern times, the wolf occurs mostly in wilderness and remote areas. The wolf can be found between sea level and {{convert|3000|m|feet|abbr=on}}. Wolves live in forests, inland [[wetland]]s, [[shrubland]]s, [[grassland]]s (including Arctic [[tundra]]), [[pasture]]s, deserts, and rocky peaks on mountains.<ref name="iucn status 2 June 2024" /> Habitat use by wolves depends on the abundance of prey, snow conditions, livestock densities, road densities, human presence and [[topography]].<ref name=Paquet2003/>

=== Diet ===

[[File:Wolf with Caribou Hindquarter.jpg|thumb|upright|left|alt=Photograph of a wolf carrying a caribou leg in its mouth|A wolf carrying a [[Reindeer|caribou]] hindquarter, [[Denali National Park]], [[Alaska]]]]

Like all land mammals that are [[pack hunter]]s, the wolf feeds predominantly on [[ungulates]] that can be divided into large size {{cvt|240|–|650|kg}} and medium size {{cvt|23|–|130|kg}}, and have a body mass similar to that of the combined mass of the [[Pack (canine)|pack]] members.<ref name=Earle1987/><ref name=Sorkin2008/> The wolf specializes in preying on the vulnerable individuals of large prey,<ref name=Paquet2003/> with a pack of 15 able to bring down an adult [[moose]].<ref name=Mech1966/> The variation in diet between wolves living on different continents is based on the variety of hoofed mammals and of available smaller and domesticated prey.<ref name=Newsome2016/>

In North America, the wolf's diet is dominated by wild large hoofed mammals (ungulates) and medium-sized mammals. In Asia and Europe, their diet is dominated by wild medium-sized hoofed mammals and domestic species. The wolf depends on wild species, and if these are not readily available, as in Asia, the wolf is more reliant on domestic species.<ref name=Newsome2016/> Across Eurasia, wolves prey mostly on [[moose]], [[red deer]], [[Capreolus|roe deer]] and [[wild boar]].{{sfn|Mech|Boitani|2003|p=107}} In North America, important range-wide prey are [[elk]], moose, [[Reindeer|caribou]], [[white-tailed deer]] and [[mule deer]].{{sfn|Mech|Boitani|2003|pp=109–110}} Prior to their extirpation from North America, [[wild horse]]s were among the most frequently consumed prey of North American wolves.<ref>{{Cite journal |last1=Landry |first1=Zoe |last2=Kim |first2=Sora |last3=Trayler |first3=Robin B. |last4=Gilbert |first4=Marisa |last5=Zazula |first5=Grant |last6=Southon |first6=John |last7=Fraser |first7=Danielle |date=1 June 2021 |title=Dietary reconstruction and evidence of prey shifting in Pleistocene and recent gray wolves (Canis lupus) from Yukon Territory |url=https://linkinghub.elsevier.com/retrieve/pii/S003101822100153X |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |language=en |volume=571 |pages=110368 |doi=10.1016/j.palaeo.2021.110368 |bibcode=2021PPP...57110368L |access-date=23 April 2024 |via=Elsevier Science Direct |issn=0031-0182|url-access=subscription }}</ref> Wolves can digest their meal in a few hours and can feed several times in one day, making quick use of large quantities of meat.{{sfn|Mech|1981|p=172}} A well-fed wolf stores fat under the skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter.{{sfn|Mech|Boitani|2003|p=201}}

Nonetheless, wolves are not fussy eaters. Smaller-sized animals that may supplement their diet include [[rodent]]s, [[hare]]s, [[insectivore]]s and smaller carnivores. They frequently eat [[waterfowl]] and their eggs. When such foods are insufficient, they prey on [[lizard]]s, [[snake]]s, and [[frog]]s, when available,{{sfn|Heptner|Naumov|1998|pp=213–231}} and have even been known to feed on [[Grasshopper|grasshoppers]].<ref>{{Cite journal |last=Barton |first=Brandon T. |last2=Hill |first2=JoVonn G. |last3=Wolff |first3=Carter L. |last4=Newsome |first4=Thomas M. |last5=Ripple |first5=William J. |last6=Lashley |first6=Marcus A. |date=18 September 2019 |title=Grasshopper consumption by grey wolves and implications for ecosystems |url=https://esajournals.onlinelibrary.wiley.com/doi/10.1002/ecy.2892 |journal=[[Ecology (journal)|Ecology]] |language=en |volume=101 |issue=2 |doi=10.1002/ecy.2892 |issn=0012-9658 |access-date=20 April 2025 |via=Wiley Online Library|url-access=subscription }}</ref> Wolves in some areas may consume fish and even marine life.<ref name="Gable2018" /><ref name="Woodford2019" /><ref name="McAllister2007" /> Wolves also consume some plant material. In Europe, they eat apples, pears, [[Common fig|figs]], melons, [[Berry|berries]] and [[Cherry|cherries]]. In North America, wolves eat [[Blueberry|blueberries]] and [[Raspberry|raspberries]]. They also eat grass, which may provide some vitamins, but is most likely used mainly to induce vomiting to rid themselves of intestinal parasites or long guard hairs.<ref name="Fuller2019" /> They are known to eat the berries of [[Sorbus|mountain-ash]], [[lily of the valley]], [[Bilberry|bilberries]], [[Vaccinium vitis-idaea|cowberries]], [[Solanum nigrum|European black nightshade]], grain crops, and the shoots of reeds.{{sfn|Heptner|Naumov|1998|pp=213–231}}

In times of scarcity, wolves will readily eat [[carrion]].{{sfn|Heptner|Naumov|1998|pp=213–231}} In Eurasian areas with dense human activity, many wolf populations are forced to subsist largely on livestock and garbage.{{sfn|Mech|Boitani|2003|p=107}} As prey in North America continue to occupy suitable habitats with low human density, North American wolves eat livestock and garbage only in dire circumstances.{{sfn|Mech|Boitani|2003|p=109}} [[Cannibalism]] is not uncommon in wolves during harsh winters, when packs often attack weak or injured wolves and may eat the bodies of dead pack members.{{sfn|Heptner|Naumov|1998|pp=213–231}}{{sfn|Mech|1981|p=180}}<ref name=Klein1995/>

=== Interactions with other predators ===

Wolves typically dominate other canid species in areas where they both occur. In North America, incidents of wolves killing coyotes are common, particularly in winter, when coyotes feed on wolf kills. Wolves may attack coyote den sites, digging out and killing their pups, though rarely eating them. There are no records of coyotes killing wolves, though coyotes may chase wolves if they outnumber them.{{sfn|Mech|Boitani|2003|pp=266–268}} According to a press release by the U.S. Department of Agriculture in 1921, the infamous [[Custer Wolf]] relied on coyotes to accompany him and warn him of danger. Though they fed from his kills, he never allowed them to approach him.<ref name=Merrit1921/> Interactions have been observed in Eurasia between wolves and golden jackals, the latter's numbers being comparatively small in areas with high wolf densities.{{sfn|Heptner|Naumov|1998|pp=164–270}}{{sfn|Mech|Boitani|2003|pp=266–268}}<ref name=Giannatos2004/> Wolves also kill [[Red fox|red]], [[Arctic fox|Arctic]] and [[corsac fox]]es, usually in disputes over carcasses, sometimes eating them.{{sfn|Heptner|Naumov|1998|pp=164–270}}{{sfn|Mech|Boitani|2003|p=269}}
[[File:Wolves bear ravens.png|thumb|350px|alt=Photograph of a wolf, a bear, coyotes and ravens competing over a kill|A wolf, a bear, coyotes and ravens compete over a kill]]

[[Brown bear]]s typically dominate wolf packs in disputes over carcasses, while wolf packs mostly prevail against bears when defending their den sites. Both species kill each other's young. Wolves eat the brown bears they kill, while brown bears seem to eat only young wolves.{{sfn|Mech|Boitani|2003|pp=261–263}} Wolf interactions with [[American black bear]]s are much rarer because of differences in habitat preferences. Wolves have been recorded on numerous occasions actively seeking out American black bears in their dens and killing them without eating them. Unlike brown bears, American black bears frequently lose against wolves in disputes over kills.{{sfn|Mech|Boitani|2003|pp=263–264}} Wolves also dominate and sometimes kill [[wolverine]]s, and will chase off those that attempt to scavenge from their kills. Wolverines escape from wolves in caves or up trees.{{sfn|Mech|Boitani|2003|p=266}}

Wolves may interact and compete with [[Felidae|felids]], such as the [[Eurasian lynx]], which may feed on smaller prey where wolves are present{{sfn|Mech|Boitani|2003|p=265}} and may be suppressed by large wolf populations.<ref name=Sunquist2002/> Wolves encounter [[cougar]]s along portions of the Rocky Mountains and adjacent mountain ranges. Wolves and cougars typically avoid encountering each other by hunting at different elevations for different prey ([[niche partitioning]]). This is more difficult during winter. Wolves in packs usually dominate cougars and can steal their kills or even kill them,{{sfn|Mech|Boitani|2003|pp=264–265}} while one-to-one encounters tend to be dominated by the cat, who likewise will kill wolves.<ref name=Jimenez2008/> Wolves more broadly affect cougar population dynamics and distribution by dominating territory and prey opportunities and disrupting the feline's behaviour.<ref name=Elbroch2015/> Wolf and [[Siberian tiger]] interactions are well-documented in the [[Russian Far East]], where tigers significantly depress wolf numbers, sometimes to the point of [[Local extinction|localized extinction]].<ref name=Miquelle2005/>{{sfn|Mech|Boitani|2003|p=265}}

In Israel, Palestine, Central Asia and India wolves may encounter [[striped hyena]]s, usually in disputes over carcasses. Striped hyenas feed extensively on wolf-killed carcasses in areas where the two species interact. One-to-one, hyenas dominate wolves, and may prey on them,<ref name=Monchot2010/> but wolf packs can drive off single or outnumbered hyenas.<ref name=Mills1998/><ref name=Nayak2015/> There is at least one case in Israel of a hyena associating and cooperating with a wolf pack.<ref name=Dinets2016/>

===Infections===
{{main|Parasites and pathogens of wolves}}
[[File:Wild Wolf Afflicted with Mange.jpg|thumb|alt=Photograph of a wolf with mange eating at a kill|Wolf with mange [[Yellowstone National Park]]]]
[[Viral disease]]s carried by wolves include: [[rabies]], [[canine distemper]], [[canine parvovirus]], [[infectious canine hepatitis]], [[papillomatosis]], and [[canine coronavirus]]. In wolves, the [[incubation period]] for rabies is eight to 21 days, and results in the host becoming agitated, deserting its pack, and travelling up to {{convert|80|km|mi|abbr=on|sigfig=1}} a day, thus increasing the risk of infecting other wolves. Although canine distemper is lethal in dogs, it has not been recorded to kill wolves, except in Canada and Alaska. The canine parvovirus, which causes death by [[dehydration]], [[electrolyte imbalance]], and [[Endotoxin|endotoxic]] shock or [[sepsis]], is largely survivable in wolves, but can be lethal to pups.{{sfn|Mech|Boitani|2003|pp=208–211}} [[Bacterial diseases]] carried by wolves include: [[brucellosis]], [[Lyme disease]], [[leptospirosis]], [[tularemia]], [[bovine tuberculosis]],{{sfn|Mech|Boitani|2003|pp=211–213}} [[listeriosis]] and [[anthrax]].{{sfn|Graves|2007|pp=77–85}} Although lyme disease can debilitate individual wolves, it does not appear to significantly affect wolf populations. Leptospirosis can be contracted through contact with infected prey or urine, and can cause [[fever]], [[Anorexia (symptom)|anorexia]], vomiting, [[anemia]], [[hematuria]], [[Jaundice|icterus]], and death.{{sfn|Mech|Boitani|2003|pp=211–213}}

Wolves are often infested with a variety of [[arthropod]] exoparasites, including [[flea]]s, [[tick]]s, [[Louse|lice]], and [[mite]]s. The most harmful to wolves, particularly pups, is the mange mite (''[[Sarcoptes scabiei]]''),{{sfn|Mech|Boitani|2003|pp=202–208}} though they rarely develop full-blown [[mange]], unlike foxes.{{sfn|Heptner|Naumov|1998|pp=164–270}} Endoparasites known to infect wolves include: [[protozoan]]s and [[helminth]]s ([[Trematoda|flukes]], [[tapeworm]]s, [[roundworm]]s and [[thorny-headed worm]]s). Most fluke species reside in the wolf's intestines. Tapeworms are commonly found in wolves, which they get though their prey, and generally cause little harm in wolves, though this depends on the number and size of the parasites, and the sensitivity of the host. Symptoms often include [[constipation]], toxic and [[allergic reaction]]s, irritation of the [[intestinal mucosa]], and [[malnutrition]]. Wolves can carry over 30 roundworm species, though most roundworm infections appear benign, depending on the number of worms and the age of the host.{{sfn|Mech|Boitani|2003|pp=202–208}}

== Behaviour ==

{{anchor|Behaviour}}
{{See also|Dog behaviour}}

=== Social structure ===

{{See also|Pack (canine)#Pack behavior in grey wolves}}
[[File:Canis lupus pallipes Mysore Zoo 2.jpg|thumb|alt=Photograph of three wolves running and biting each other|Indian wolves at the [[Mysore Zoo]]]]

The wolf is a [[Sociality|social animal]].{{sfn|Heptner|Naumov|1998|pp=164–270}} Its populations consist of packs and lone wolves, most lone wolves being temporarily alone while they disperse from packs to form their own or join another one.{{sfn|Mech|Boitani|2003|p=164}} The wolf's basic social unit is the [[nuclear family]] consisting of a [[mated pair]] accompanied by their offspring.{{sfn|Heptner|Naumov|1998|pp=164–270}} The average pack size in North America is eight wolves and 5.5 in Europe.<ref name=Miklosi2015/> The average pack across Eurasia consists of a family of eight wolves (two adults, juveniles, and yearlings),{{sfn|Heptner|Naumov|1998|pp=164–270}} or sometimes two or three such families,<ref name=Paquet2003/> with examples of exceptionally large packs consisting of up to 42 wolves being known.{{sfn|Mech|Boitani|2003|pp=2–3, 28}} [[Cortisol]] levels in wolves rise significantly when a pack member dies, indicating the presence of stress.<ref name=Molnar2015/> During times of prey abundance caused by calving or migration, different wolf packs may join together temporarily.{{sfn|Heptner|Naumov|1998|pp=164–270}}

Offspring typically stay in the pack for 10–54 months before dispersing.{{sfn|Mech|Boitani|2003|pp=1–2}} Triggers for dispersal include the onset of [[sexual maturity]] and competition within the pack for food.{{sfn|Mech|Boitani|2003|pp=12–13}} The distance travelled by dispersing wolves varies widely; some stay in the vicinity of the parental group, while other individuals may travel great distances of upwards of {{convert|206|km|mi|abbr=on}}, {{convert|390|km|mi|abbr=on}}, and {{convert|670|km|mi|abbr=on}} from their natal (birth) packs.<ref name=Nowak1983/> A new pack is usually founded by an unrelated dispersing male and female, travelling together in search of an area devoid of other hostile packs.{{sfn|Mech|Boitani|2003|p=38}} Wolf packs rarely adopt other wolves into their fold and typically kill them. In the rare cases where other wolves are adopted, the adoptee is almost invariably an immature animal of one to three years old, and unlikely to compete for breeding rights with the mated pair. This usually occurs between the months of February and May. Adopted males may mate with an available pack female and then form their own pack. In some cases, a lone wolf is adopted into a pack to replace a deceased breeder.{{sfn|Mech|Boitani|2003|pp=2–3, 28}}

Wolves are [[Territory (animal)|territorial]] and generally establish territories far larger than they require to survive assuring a steady supply of prey. Territory size depends largely on the amount of prey available and the age of the pack's pups. They tend to increase in size in areas with low prey populations,<ref name=Jedrzejewski2007/> or when the pups reach the age of six months when they have the same nutritional needs as adults.{{sfn|Mech|Boitani|2003|pp=19–26}} Wolf packs travel constantly in search of prey, covering roughly 9% of their territory per day, on average {{cvt|25|km/d}}. The core of their territory is on average {{cvt|35|km2}} where they spend 50% of their time.<ref name=Jedrzejewski2007/> Prey density tends to be much higher on the territory's periphery. Wolves tend to avoid hunting on the fringes of their range to avoid fatal confrontations with neighbouring packs.<ref name=Mech1977/> The smallest territory on record was held by a pack of six wolves in northeastern Minnesota, which occupied an estimated {{cvt|33|km2}}, while the largest was held by an Alaskan pack of ten wolves encompassing {{cvt|6,272|km2}}.{{sfn|Mech|Boitani|2003|pp=19–26}} Wolf packs are typically settled, and usually leave their accustomed ranges only during severe food shortages.{{sfn|Heptner|Naumov|1998|pp=164–270}} Territorial fights are among the principal causes of wolf mortality, one study concluding that 14–65% of wolf deaths in Minnesota and the [[Denali National Park and Preserve]] were due to other wolves.<ref name=Mech2003/>

===Communication===
{{main article|Wolf communication}}
{{listen
| filename  = Wolf howls.ogg
| title = Wolves howling
| format = [[Ogg]]
| filename2 = rallying.ogg
| title2 = Rallying cry
| format2 = [[Ogg]]
}}
Wolves communicate using vocalizations, body postures, scent, touch, and taste.{{sfn|Mech|Boitani|2003|pp=66–103}} The phases of the moon have no effect on wolf vocalization, and despite popular belief, wolves do not howl at the Moon.{{sfn|Busch|2007|p=59}} Wolves [[howling|howl]] to assemble the pack usually before and after hunts, to pass on an alarm particularly at a den site, to locate each other during a storm, while crossing unfamiliar territory, and to communicate across great distances.{{sfn|Lopez|1978|p=38}} Wolf howls can under certain conditions be heard over areas of up to {{cvt|130|km2}}.<ref name=Paquet2003/> Other vocalizations include [[growling|growls]], [[bark (sound)|barks]] and whines. Wolves do not bark as loudly or continuously as dogs do in confrontations, rather barking a few times and then retreating from a perceived danger.{{sfn|Lopez|1978|pp=39–41}} Aggressive or self-assertive wolves are characterized by their slow and deliberate movements, high body [[Posture (psychology)|posture]] and raised [[hackles]], while submissive ones carry their bodies low, flatten their fur, and lower their ears and tail.{{sfn|Mech|Boitani|2003|p=90}}

Scent marking involves urine, feces, and [[preputial gland|preputial]] and anal gland scents. This is more effective at advertising territory than howling and is often used in combination with scratch marks. Wolves increase their rate of scent marking when they encounter the marks of wolves from other packs. Lone wolves will rarely mark, but newly bonded pairs will scent mark the most.<ref name=Paquet2003/> These marks are generally left every {{cvt|240|m}} throughout the territory on regular travelways and junctions. Such markers can last for two to three weeks,{{sfn|Mech|Boitani|2003|pp=19–26}} and are typically placed near rocks, boulders, trees, or the skeletons of large animals.{{sfn|Heptner|Naumov|1998|pp=164–270}} Raised leg [[urination]] is considered to be one of the most important forms of scent communication in the wolf, making up 60–80% of all scent marks observed.<ref name=Peters1975/>

=== Reproduction{{anchor|Reproduction_and_development}} ===

{{See also|Canine reproduction}}
[[File:Korean wolves mating (cropped).jpg|left|thumb|alt=Photograph of a pair of mating wolves|Korean wolves mating in the [[Tama Zoological Park]], Japan]]
Wolves are [[Monogamy in animals|monogamous]], mated pairs usually remaining together for life. Should one of the pair die, another mate is found quickly.{{sfn|Heptner|Naumov|1998|p=248}} With wolves in the wild, inbreeding does not occur where outbreeding is possible.{{sfn|Mech|Boitani|2003|p=5}} Wolves become mature at the age of two years and sexually mature from the age of three years.{{sfn|Heptner|Naumov|1998|p=248}} The age of first breeding in wolves depends largely on environmental factors: when food is plentiful, or when wolf populations are heavily managed, wolves can rear pups at younger ages to better exploit abundant resources. Females are capable of producing pups every year, one [[Litter (zoology)|litter]] annually being the average.{{sfn|Mech|Boitani|2003|p=175}} [[Oestrus]] and [[Rut (mammalian reproduction)|rut]] begin in the second half of winter and lasts for two weeks.{{sfn|Heptner|Naumov|1998|p=248}}
[[File:Iberian Wolf 01.jpg|thumb|alt=Photograph of wolf pups stimulating their mother to regurgitate some food|[[Iberian wolf]] pups stimulating their mother to regurgitate some food]]

Dens are usually constructed for pups during the summer period. When building dens, females make use of natural shelters like fissures in rocks, cliffs overhanging riverbanks and holes thickly covered by vegetation. Sometimes, the den is the appropriated burrow of smaller animals such as foxes, badgers or marmots. An appropriated den is often widened and partly remade. On rare occasions, female wolves dig burrows themselves, which are usually small and short with one to three openings. The den is usually constructed not more than {{cvt|500|m}} away from a water source. It typically faces southwards where it can be better warmed by sunlight exposure, and the snow can thaw more quickly. Resting places, play areas for the pups, and food remains are commonly found around wolf dens. The odor of urine and rotting food emanating from the denning area often attracts scavenging birds like [[magpie]]s and [[raven]]s. Though they mostly avoid areas within human sight, wolves have been known to nest near homes, paved [[road]]s and [[railway]]s.{{sfn|Heptner|Naumov|1998|pp=234–238}} During pregnancy, female wolves remain in a den located away from the peripheral zone of their territories, where violent encounters with other packs are less likely to occur.{{sfn|Mech|Boitani|2003|pp=42–46}}

The [[gestation period]] lasts 62–75 days with pups usually being born in the spring months or early summer in very cold places such as on the tundra. Young females give birth to four to five young, and older females from six to eight young and up to 14. Their mortality rate is 60–80%.{{sfn|Heptner|Naumov|1998|pp=249–254}} Newborn wolf pups look similar to [[German Shepherd|German Shepherd Dog]] pups.{{sfn|Mech|Boitani|2003|p=47}} They are born blind and deaf and are covered in short soft greyish-brown fur. They weigh {{cvt|300|–|500|g}} at birth and begin to see after nine to 12 days. The milk canines erupt after one month. Pups first leave the den after three weeks. At one-and-a-half months of age, they are agile enough to flee from danger. Mother wolves do not leave the den for the first few weeks, relying on the fathers to provide food for them and their young. Pups begin to eat solid food at the age of three to four weeks. They have a fast growth rate during their first four months of life: during this period, a pup's weight can increase nearly 30 times.{{sfn|Heptner|Naumov|1998|pp=249–254}}{{sfn|Mech|Boitani|2003|pp=46–49}} Wolf pups begin play-fighting at the age of three weeks, though unlike young coyotes and foxes, their bites are gentle and controlled. Actual fights to establish hierarchy usually occur at five to eight weeks of age. This is in contrast to young coyotes and foxes, which may begin fighting even before the onset of play behaviour.<ref name=Fox1978/> By autumn, the pups are mature enough to accompany the adults on hunts for large prey.{{sfn|Mech|Boitani|2003|pp=42–46}}

===Hunting and feeding===
{{main article|Hunting behavior of gray wolves}}
[[File:Wolves and elk (cropped).jpg|thumb|right|alt=Aerial photograph a bull elk in winter being pursued by four wolves |Wolves pursuing a bull elk]]
Single wolves or mated pairs typically have higher success rates in hunting than do large packs; single wolves have occasionally been observed to kill large prey such as moose, [[bison]] and [[muskox]]en unaided.{{sfn|Mech|Boitani|2003|pp=119–121}}<ref name=Thurber1993/> The size of a wolf hunting pack is related to the number of pups that survived the previous winter, adult survival, and the rate of dispersing wolves leaving the pack. The optimal pack size for hunting elk is four wolves, and for bison a large pack size is more successful.{{sfn|Mech|Smith|MacNulty|2015|p=4}} Wolves move around their territory when hunting, using the same trails for extended periods.{{sfn|Heptner|Naumov|1998|p=233}} Wolves are nocturnal predators. During the winter, a pack will commence hunting in the twilight of early evening and will hunt all night, traveling tens of kilometres. Sometimes hunting large prey occurs during the day. During the summer, wolves generally tend to hunt individually, ambushing their prey and rarely giving pursuit.{{sfn|Heptner|Naumov|1998|p=240}} 

When hunting large gregarious prey, wolves will try to isolate an individual from its group.<ref name=MacNulty2007/> If successful, a wolf pack can bring down game that will feed it for days, but one error in judgement can lead to serious injury or death. Most large prey have developed defensive adaptations and behaviours. Wolves have been killed while attempting to bring down bison, elk, moose, muskoxen, and even by one of their smallest hoofed prey, the white-tailed deer. With smaller prey like [[beaver]], geese, and hares, there is no risk to the wolf.{{sfn|Mech|Smith|MacNulty|2015|pp=1–3}} Although people often believe wolves can easily overcome any of their prey, their success rate in hunting hoofed prey is usually low.{{sfn|Mech|Smith|MacNulty|2015|p=7}} 
[[File:Wolves eating deer.jpeg|thumb|right|alt=Photograph of two wolves eating a deer carcass at night|Two wolves feeding on a [[white-tailed deer]]]]
The wolf must give chase and gain on its fleeing prey, slow it down by biting through thick hair and hide, and then disable it enough to begin feeding.{{sfn|Mech|Smith|MacNulty|2015|pp=1–3}} Wolves may wound large prey and then lie around resting for hours before killing it when it is weaker due to blood loss, thereby lessening the risk of injury to themselves.{{sfn|Mech|Smith|MacNulty|2015|p=82–89}} With medium-sized prey, such as roe deer or [[sheep]], wolves kill by [[Throat clamp|biting the throat]], severing nerve tracks and the [[carotid artery]], thus causing the animal to die within a few seconds to a minute. With small, [[mouse]]like prey, wolves leap in a high arc and immobilize it with their forepaws.<ref name=Zimen/>

Once prey is brought down, wolves begin to feed excitedly, ripping and tugging at the carcass in all directions, and bolting down large chunks of it.{{sfn|Mech|1981|p=185}} The breeding pair typically monopolizes food to continue producing pups. When food is scarce, this is done at the expense of other family members, especially non-pups.{{sfn|Mech|Boitani|2003|p=58}} Wolves typically commence feeding by gorging on the larger internal organs, like the [[heart]], [[liver]], [[lung]]s, and [[stomach]] lining. The [[kidney]]s and [[spleen]] are eaten once they are exposed, followed by the muscles.{{sfn|Mech|Boitani|2003|pp=122–125}} A wolf can eat 15–19% of its body weight in one sitting.{{sfn|Mech|Boitani|2003|p=201}}

== Status and conservation ==
{{Further|List of gray wolf populations by country}}
<!-- Please do not expand. This subject already has its own article -->
The global wild wolf population in 2003 was estimated at 300,000.{{sfn|Mech|Boitani|2003|p=230}} Wolf population declines have been arrested since the 1970s. This has fostered recolonization and reintroduction in parts of its former range as a result of legal protection, changes in land use, and rural human population shifts to cities. Competition with humans for livestock and game species, concerns over the danger posed by wolves to people, and habitat fragmentation pose a continued threat to the wolf. Despite these threats, the [[International Union for Conservation of Nature|IUCN]] classifies the wolf as [[Least Concern]] on its [[IUCN Red List|Red List]] due to its relatively widespread range and stable population. The species is listed under [[CITES#Appendix II|Appendix{{spaces}}II]] of the [[Convention on International Trade in Endangered Species of Wild Fauna and Flora]] (CITES), meaning international trade in the species (including parts and derivatives) is regulated. However, populations of [[Bhutan]], [[India]], [[Nepal]] and [[Pakistan]] are listed in [[CITES#Appendix I|Appendix{{spaces}}I]] which prohibits commercial international trade in wild-sourced specimens.<ref name="iucn status 2 June 2024" />

=== North America ===
<!-- Please do not expand. This subject already has its own article -->
[[File:Mexican Wolf 2 yfb-edit 1.jpg|thumb|right|alt=Photograph of a wolf running on a grassy plain with enclosing fence in background|Captive Mexican wolf at [[Sevilleta National Wildlife Refuge]] in New Mexico, as part of [[Wolf reintroduction|reintroduction]]]]

In Canada, 50,000–60,000 wolves live in 80% of their historical range, making Canada an important stronghold for the species.<ref name=Paquet2003/> Under Canadian law, [[First Nations in Canada|First Nations]] people can hunt wolves without restrictions, but others must acquire licenses for the hunting and trapping seasons. As many as 4,000 wolves may be harvested in Canada each year.{{sfn|Mech|Boitani|2003|pp=321–324}} The wolf is a protected species in national parks under the [[Canada National Parks Act]].<ref name=Justice2019/> In Alaska, 7,000–11,000 wolves are found on 85% of the state's {{cvt|1,517,733|km2}} area. Wolves may be hunted or trapped with a license; around 1,200 wolves are harvested annually.<ref name=Alaska2019/>

In the [[contiguous United States]], wolf declines were caused by the expansion of agriculture, the decimation of the wolf's main prey species like the American bison, and extermination campaigns.<ref name=Paquet2003/> Wolves were given protection under the [[Endangered Species Act]] (ESA) of 1973, and have since returned to parts of their former range thanks to both natural recolonizations and [[History of wolves in Yellowstone#Re-introduction (1995–present)|reintroductions in Yellowstone and Idaho]].<ref name=FWS2007/> The [[repopulation of wolves in Midwestern United States]] has been concentrated in the [[Western Great Lakes forests|Great Lakes]] states of Minnesota, Wisconsin and Michigan where wolves number over 4,000 as of 2018.<ref name=USFWGreatLakes/> Wolves also occupy much of the northern Rocky Mountains region and the northwest, with a total population over 3,000 as of the 2020s.<ref name=WildWolfUS/> In Mexico and parts of the southwestern United States, the Mexican and U.S. governments collaborated from 1977 to 1980 in capturing all Mexican wolves remaining in the wild to prevent their extinction and established captive breeding programs for reintroduction.<ref name=Nie2003/> As of 2024, the reintroduced [[Mexican wolf]] population numbers over 250 individuals.<ref name=Mexicanwolf/>

=== Eurasia ===
<!-- Please do not expand. This subject already has its own article -->
[[File:Wolf - populations in Europe.png|thumb|right|alt=Map showing the wolf's range in Europe and surrounding areas]]<!--[[wp:caption]] "Not every image ..."-->

The [[European Union]] has 20,300 wolves with breeding packs in 23 countries.<ref name=EUcomm2023/> In many EU countries, the wolf is strictly protected under the 1979 [[Berne Convention on the Conservation of European Wildlife and Natural Habitats]] (Appendix{{spaces}}II) and the 1992 [[Habitats Directive|Council Directive 92/43/EEC on the Conservation of Natural Habitats and of Wild Fauna and Flora]] (Annex{{spaces}}II and{{spaces}}IV). There is extensive legal protection in many European countries, although there are national exceptions.<ref name="iucn status 2 June 2024" /><ref>[[European Commission]]: [http://ec.europa.eu/environment/nature/conservation/species/carnivores/pdf/task_1_part1_statusoflcineurope.pdf ''Status, management and distribution of large carnivores—bear, lynx, wolf & wolverine—in Europe December 2012.''] page 50.</ref>

Wolves have been persecuted in Europe for centuries, having been exterminated in [[Wolves in Great Britain|Great Britain]] by 1684, in [[Wolves in Ireland|Ireland]] by 1770, in Central Europe by 1899, in France by the 1930s, and in much of Scandinavia by the early 1970s. They continued to survive in parts of Finland, Eastern Europe and Southern Europe.{{sfn|Mech|Boitani|2003|pp=318–320}} Since 1980, European wolves have rebounded and expanded into parts of their former range. The decline of the traditional pastoral and rural economies seems to have ended the need to exterminate the wolf in parts of Europe.{{sfn|Mech|Boitani|2003|pp=321–324}} As of 2016, estimates of wolf numbers include: 4,000 in the Balkans, 3,460–3,849 in the [[Carpathian Mountains]], 1,700–2,240 in the [[Baltic states]], 1,100–2,400 in the [[Italian Peninsula]], and around 2,500 in the northwest [[Iberian peninsula]] as of 2007.<ref name=EUcomm2019/> In a study of wolf conservation in Sweden, it was found that there was little  opposition between the policies of the European Union and those of the Swedish officials implementing domestic policy.<ref>''The Wolf Dilemma: Following the Practices of Several Actors in Swedish Large Carnivore Management.'' Ramsey, Morag (2015) https://www.diva-portal.org/smash/get/diva2:821120/FULLTEXT01.pdf Retrieved 30 September 2023</ref>

In the [[former Soviet Union]], wolf populations have retained much of their historical range despite Soviet-era large scale extermination campaigns. Their numbers range from 1,500 in Georgia, to 20,000 in Kazakhstan and up to 45,000 in Russia.<ref name=Goldthorpe2016/> In Russia, the wolf is regarded as a pest because of its attacks on livestock, and wolf management means controlling their numbers by destroying them throughout the year. Russian history over the past century shows that reduced hunting leads to an abundance of wolves.<ref name=Baskin2016/> The Russian government has continued to pay bounties for wolves and annual harvests of 20–30% do not appear to significantly affect their numbers.<ref name=Russia/>
[[File:Desert wolf 1.jpg|thumb|left|upright|alt=Image of a wolf at night with glowing eyes|A wolf in southern [[Israel]]]]

In the Middle East, only Israel and Oman give wolves explicit legal protection.<ref name=Fisher2019/> Israel has protected its wolves since 1954 and has maintained a moderately sized population of 150 through effective enforcement of conservation policies. These wolves have moved into neighboring countries. Approximately 300–600 wolves inhabit the [[Arabian Peninsula]].{{sfn|Mech|Boitani|2003|pp=323, 327}} The wolf also appears to be widespread in Iran.{{sfn|Busch|2007|p=231}} Turkey has an estimated population of about 7,000 wolves.<ref name=Sekercioglu2013/> Outside of Turkey, wolf populations in the Middle East may total 1,000–2,000.<ref name=Fisher2019/>

In southern Asia, the northern regions of Afghanistan and [[Pakistan]] are important strongholds for wolves. The wolf has been protected in [[India]] since 1972.{{sfn|Mech|Boitani|2003|p=327}} The Indian wolf is distributed across the states of [[Gujarat]], [[Rajasthan]], [[Haryana]], [[Uttar Pradesh]], [[Madhya Pradesh]], [[Maharashtra]], [[Karnataka]] and [[Andhra Pradesh]].<ref name=Yadvendradev/> As of 2019, it is estimated that there are around 2,000–3,000 Indian wolves in the country.<ref name=Sharma/> In East Asia, Mongolia's population numbers 10,000–20,000. In China, [[Heilongjiang]] has roughly 650 wolves, [[Xinjiang]] has 10,000 and [[Tibet Autonomous Region|Tibet]] has 2,000.{{sfn|Mech|Boitani|2003|p=328}} 2017 evidence suggests that wolves range across all of mainland China.<ref name=Larson2017/> Wolves have been historically persecuted in China{{sfn|Busch|2007|p=232}} but have been legally protected since 1998.<ref name=Xu2015/> The last [[Japanese wolf]] was captured and killed in 1905.<ref name=Ishiguro2009/>

== Relationships with humans ==
=== In culture ===
==== In folklore, religion and mythology ====
<!-- Please do not add any more examples to this section. This subject already has its own article -->
{{Main|Wolves in folklore, religion and mythology}}
{{See also|Wolves in heraldry}}
[[File:Lupa Capitolina, Rome.jpg|thumb|left|alt=Photograph of the sculpture ''Capitoline Wolf'' showing of the mythical she-wolf feeding the twins Romulus and Remus|The ''[[Capitoline Wolf]]'', sculpture of the mythical [[She-wolf (Roman mythology)|she-wolf]] feeding the twins [[Romulus and Remus]], from the legend of the [[founding of Rome]], Italy, 13th century AD. (The twins are a 15th-century addition.)]]

The wolf is a common motif in the mythologies and cosmologies of peoples throughout its historical range. The [[Ancient Greeks]] associated wolves with [[Apollo]], the god of light and order.{{sfn|Mech|Boitani|2003|p=292}} The [[Ancient Romans]] connected the wolf with their god of war and agriculture [[Mars (mythology)|Mars]],{{sfn|Lopez|1978|p=210}} and believed their city's founders, [[Romulus and Remus]], were suckled by a [[She-wolf (Roman mythology)|she-wolf]].{{sfn|Marvin|2012|p=128}} [[Norse mythology]] includes the feared giant wolf [[Fenrir]],<ref name=Symbolism/> and [[Geri and Freki]], [[Odin]]'s faithful pets.{{sfn|Marvin|2012|p=78}}

In [[Chinese astronomy]], the wolf represents [[Sirius]] and guards the heavenly gate. In China, the wolf was traditionally associated with greed and cruelty and wolf epithets were used to describe negative behaviours such as cruelty ("wolf's heart"), mistrust ("wolf's look") and lechery ("wolf-sex"). In both [[Hinduism]] and [[Buddhism]], the wolf is ridden by gods of protection. In [[Vedic]] Hinduism, the wolf is a symbol of the night and the daytime [[quail]] must escape from its jaws. In [[Tantric Buddhism]], wolves are depicted as inhabitants of graveyards and destroyers of corpses.<ref name=Symbolism/>

In the [[Pawnee people|Pawnee]] creation myth, the wolf was the first animal brought to Earth. When humans killed it, they were punished with death, destruction and the loss of immortality.{{sfn|Lopez|1978|p=133}} For the Pawnee, Sirius is the "wolf star" and its disappearance and reappearance signified the wolf moving to and from the spirit world. Both Pawnee and [[Blackfoot]] call the [[Milky Way]] the "wolf trail".{{sfn|Busch|2007|p=110}} The wolf is also an important [[Crest (heraldry)|crest]] symbol for clans of the Pacific Northwest like the [[Kwakwakaʼwakw]].<ref name=Symbolism/>

The concept of people turning into wolves, and the inverse, has been present in many cultures. One [[Greek myth]] tells of [[Lycaon (king of Arcadia)|Lycaon]] being transformed into a wolf by [[Zeus]] as punishment for his evil deeds.{{sfn|Marvin|2012|p=47}} The legend of the [[werewolf]] has been widespread in [[European folklore]] and involves people willingly turning into wolves to attack and kill others.{{sfn|Marvin|2012|p=50}} The [[Navajo]] have traditionally believed that [[skin-walker|witches]] would turn into wolves by donning wolf skins and would kill people and raid graveyards.{{sfn|Lopez|1978|p=123}} The [[Dena'ina]] believed wolves were once men and viewed them as brothers.{{sfn|Mech|Boitani|2003|p=292}}

==== In fable and literature ====

{{See also|List of fictional wolves}}
[[Aesop]] featured wolves in several of his [[Aesop's Fables|fables]], playing on the concerns of Ancient Greece's settled, sheep-herding world. His most famous is the fable of "[[The Boy Who Cried Wolf]]", which is directed at those who knowingly raise false alarms, and from which the idiomatic phrase "to [[wikt:cry wolf|cry wolf]]" is derived. Some of his other fables concentrate on maintaining the trust between shepherds and guard dogs in their vigilance against wolves, as well as anxieties over the close relationship between wolves and dogs. Although Aesop used wolves to warn, criticize and moralize about human behaviour, his portrayals added to the wolf's image as a deceitful and dangerous animal. The [[Bible]] uses an image of a wolf lying with a lamb in a utopian vision of the future. In the [[New Testament]], [[Jesus]] is said to have used wolves as illustrations of the dangers his followers, whom he represents as sheep, would face should they follow him.{{sfn|Marvin|2012|pp=38–45}}
[[File:Dore ridinghood.jpg|thumb|upright|right|alt=An illustration of Red Riding Hood meeting the wolf|''[[Little Red Riding Hood]]'' (1883), [[Gustave Doré]]]] 
Isengrim the wolf, a character first appearing in the 12th-century Latin poem ''[[Ysengrimus]]'', is a major character in the [[Reynard]] Cycle, where he stands for the low nobility, whilst his adversary, Reynard the fox, represents the peasant hero. Isengrim is forever the victim of Reynard's wit and cruelty, often dying at the end of each story.{{sfn|Lopez|1978|p=259}} The tale of "[[Little Red Riding Hood]]", first written in 1697 by [[Charles Perrault]], is considered to have further contributed to the wolf's negative reputation in the Western world. The [[Big Bad Wolf]] is portrayed as a villain capable of imitating human speech and disguising itself with human clothing. The character has been interpreted as an allegorical [[sexual predator]].{{sfn|Marvin|2012|pp=64–70}} Villainous wolf characters also appear in ''[[The Three Little Pigs]]'' and "[[The Wolf and the Seven Young Goats]]".{{sfn|Lopez|1978|p=263}} The hunting of wolves, and their attacks on humans and livestock, feature prominently in [[Russian literature]], and are included in the works of [[Leo Tolstoy]], [[Anton Chekhov]], [[Nikolay Nekrasov]], [[Ivan Bunin]], [[Leonid Pavlovich Sabaneyev]], and others. Tolstoy's ''[[War and Peace]]'' and Chekhov's ''Peasants'' both feature scenes in which wolves are hunted with hounds and [[Borzoi]]s.{{sfn|Graves|2007|pp=21, 123}} The musical ''[[Peter and the Wolf]]'' involves a wolf being captured for eating a duck, but is spared and sent to a zoo.{{sfn|Marvin|2012|p=162}}

Wolves are among the central characters of [[Rudyard Kipling]]'s ''[[The Jungle Book]]''. His portrayal of wolves has been praised posthumously by wolf biologists for his depiction of them: rather than being villainous or gluttonous, as was common in wolf portrayals at the time of the book's publication, they are shown as living in amiable family groups and drawing on the experience of infirm but experienced elder pack members.<ref name=Kipling/> [[Farley Mowat]]'s largely fictional 1963 memoir ''[[Never Cry Wolf]]'' is widely considered to be the most popular book on wolves, having been adapted into a [[Never Cry Wolf (film)|Hollywood film]] and taught in several schools decades after its publication. Although credited with having changed popular perceptions on wolves by portraying them as loving, cooperative and noble, it has been criticized for its idealization of wolves and its factual inaccuracies.{{sfn|Mech|Boitani|2003|p=294}}<ref name=Jones/><ref name="Nevercrywolf"/>

=== Conflicts ===

Human presence appears to stress wolves, as seen by increased [[cortisol]] levels in instances such as snowmobiling near their territory.<ref name=Creel/>

==== Predation on livestock ====

[[File:The Allendale Wolf.jpg|thumb|left|alt=Black and white photograph of a dead wolf with "The Allendale Wolf" written on the bottom|A 1905 postcard of the [[Hexham wolf]], an escaped wolf shot for killing livestock in England]]

Livestock depredation has been one of the primary reasons for hunting wolves and can pose a severe problem for wolf conservation. As well as causing economic losses, the threat of wolf predation causes great stress on livestock producers, and no foolproof solution of preventing such attacks short of exterminating wolves has been found.{{sfn|Mech|Boitani|2003|p=305}} Some nations help offset economic losses to wolves through compensation programs or state insurance.{{sfn|Mech|Boitani|2003|p=309}} Domesticated animals are easy prey for wolves, as they have been bred under constant human protection, and are thus unable to defend themselves very well.{{sfn|Mech|1981|p=173}} Wolves typically resort to attacking livestock when wild prey is depleted.<ref>{{Cite web|last=Levy|first=Max G.|date=December 11, 2020|title=These Non-Lethal Methods Encouraged by Science Can Keep Wolves From Killing Livestock|url=https://www.smithsonianmag.com/science-nature/these-non-lethal-methods-encouraged-science-can-keep-wolves-killing-livestock-180976505/|access-date=2020-12-16|website=Smithsonian Magazine|language=en}}</ref> In Eurasia, a large part of the diet of some wolf populations consists of livestock, while such incidents are rare in North America, where healthy populations of wild prey have been largely restored.{{sfn|Mech|Boitani|2003|p=305}}

The majority of losses occur during the summer grazing period, untended livestock in remote pastures being the most vulnerable to wolf predation.{{sfn|Mech|Boitani|2003|p=307}} The most frequently targeted livestock species are sheep (Europe), [[Reindeer herding|domestic reindeer]] (northern Scandinavia), [[goat]]s (India), [[horse]]s (Mongolia), [[cattle]] and [[Domestic turkey|turkeys]] (North America).{{sfn|Mech|Boitani|2003|p=305}} The number of animals killed in single attacks varies according to species: most attacks on cattle and horses result in one death, while turkeys, sheep and domestic reindeer may be killed in surplus.{{sfn|Mech|Boitani|2003|p=306}} Wolves mainly attack livestock when the animals are grazing, though they occasionally break into fenced enclosures.{{sfn|Graves|2007|p=45}}

==== Competition with dogs ====

A review of the studies on the competitive effects of dogs on [[Sympatry|sympatric]] carnivores did not mention any research on competition between dogs and wolves.<ref name=Vanak2014/><ref name=Lescureaux2014 /> Competition would favour the wolf, which is known to kill dogs; however, wolves usually live in pairs or in small packs in areas with high human persecution, giving them a disadvantage when facing large groups of dogs.<ref name=Lescureaux2014 /><ref name=Boitani1983/>

Wolves kill dogs on occasion, and some wolf populations rely on dogs as an important food source. In Croatia, wolves kill more dogs than sheep, and wolves in Russia appear to limit stray dog populations. Wolves may display unusually bold behaviour when attacking dogs accompanied by people, sometimes ignoring nearby humans. Wolf attacks on dogs may occur both in house yards and in forests. Wolf attacks on hunting dogs are considered a major problem in Scandinavia and Wisconsin.{{sfn|Mech|Boitani|2003|p=305}}<ref name="Jess" /> Although the number of dogs killed each year by wolves is relatively low, it induces a fear of wolves entering villages and farmyards to prey on them. In many cultures, dogs are seen as family members, or at least working team members, and losing one can lead to strong emotional responses such as demanding more liberal hunting regulations.<ref name=Lescureaux2014 />

Dogs that are employed to guard sheep help to mitigate human–wolf conflicts, and are often proposed as one of the non-lethal tools in the conservation of wolves.<ref name=Lescureaux2014 /><ref name=Shivik2006/> Shepherd dogs are not particularly aggressive, but they can disrupt potential wolf predation by displaying what is to the wolf ambiguous behaviours, such as barking, social greeting, invitation to play or aggression. The historical use of shepherd dogs across Eurasia has been effective against wolf predation,<ref name=Lescureaux2014 /><ref name=Coppinger1995 /> especially when confining sheep in the presence of several livestock guardian dogs.<ref name=Lescureaux2014 /><ref name=Espuno2004/> Shepherd dogs are sometimes killed by wolves.<ref name=Lescureaux2014 />

==== Attacks on humans ====

{{Main|Wolf attack|List of wolf attacks}}
[[File:Petits Paysans surpris par un loup.jpg|thumb|alt=Painting of a wolf snarling at three children|''Country children surprised by a wolf'' (1833) by François Grenier de Saint-Martin]]

The fear of wolves has been pervasive in many societies, though humans are not part of the wolf's natural prey.<ref name="Linnell" /> How wolves react to humans depends largely on their prior experience with people: wolves lacking any negative experience of humans, or which are food-conditioned, may show little fear of people.{{sfn|Mech|Boitani|2003|pp=300–304}} Although wolves may react aggressively when provoked, such attacks are mostly limited to quick bites on extremities, and the attacks are not pressed.<ref name="Linnell"/>

Predatory attacks may be preceded by a long period of [[habituation]], in which wolves gradually lose their fear of humans. The victims are repeatedly bitten on the head and face, and are then dragged off and consumed unless the wolves are driven off. Such attacks typically occur only locally and do not stop until the wolves involved are eliminated. Predatory attacks can occur at any time of the year, with a peak in the June–August period, when the chances of people entering forested areas (for livestock [[grazing]] or berry and mushroom picking) increase.<ref name="Linnell" /> Cases of non-rabid wolf attacks in winter have been recorded in [[Belarus]], [[Kirov Oblast|Kirov]] and [[Irkutsk Oblast|Irkutsk]] oblasts, [[Karelia]] and [[Ukraine]]. Also, wolves with pups experience greater food stresses during this period.{{sfn|Heptner|Naumov|1998|pp=164–270}} The majority of victims of predatory wolf attacks are children under the age of 18 and, in the rare cases where adults are killed, the victims are almost always women.<ref name="Linnell" /> Indian wolves have a history of preying on children, a phenomenon called "child-lifting". They may be taken primarily in the spring and summer periods during the evening hours, and often within human settlements.<ref name="Rajpurohit1999"/>

Cases of rabid wolves are low when compared to other species, as wolves do not serve as primary reservoirs of the disease, but can be infected by animals such as dogs, jackals and foxes. Incidents of rabies in wolves are very rare in North America, though numerous in the eastern [[Mediterranean]], the [[Middle East]] and [[Central Asia]]. Wolves apparently develop the "furious" phase of rabies to a very high degree. This, coupled with their size and strength, makes rabid wolves perhaps the most dangerous of rabid animals.<ref name="Linnell" /> Bites from rabid wolves are 15 times more dangerous than those of rabid dogs.{{sfn|Heptner|Naumov|1998|p=267}} Rabid wolves usually act alone, travelling large distances and often biting large numbers of people and domestic animals. Most rabid wolf attacks occur in the spring and autumn periods. Unlike with predatory attacks, the victims of rabid wolves are not eaten, and the attacks generally occur only on a single day. The victims are chosen at random, though most cases involve adult men. During the fifty years up to 2002, there were eight fatal attacks in Europe and Russia, and more than two hundred in southern Asia.<ref name="Linnell" />

====Human hunting of wolves ====

{{Main|Wolf hunting|Wolf hunting with dogs}}
{{See also|Wolf hunting#Quarry|l1=Human uses of hunted wolves}}
[[File:Provedenie meropriyatii po otstrelu vrednyh givotnyh na territorii Kamyshinskogo raiona Volgogradskoi oblasti.jpg|thumb|alt=Two men with guns behind nine carcasses of hunted wolves|Carcasses of hunted wolves in [[Volgograd Oblast]], Russia]]

[[Theodore Roosevelt]] said wolves are difficult to hunt because of their elusiveness, sharp senses, high endurance, and ability to quickly incapacitate and kill hunting dogs.<ref name="Roosevelt"/> Historic methods included killing of spring-born litters in their dens, [[coursing]] with dogs (usually combinations of [[sighthound]]s, [[Bloodhound]]s and [[Fox Terrier]]s), poisoning with [[strychnine]], and [[trapping]].{{sfn|Lopez|1978|p=108}}{{sfn|Graves|2007|pp=121–140}}

A popular method of wolf hunting in Russia involves trapping a pack within a small area by encircling it with [[fladry]] poles carrying a human scent. This method relies heavily on the wolf's fear of human scents, though it can lose its effectiveness when wolves become accustomed to the odor. Some hunters can lure wolves by imitating their calls. In [[Kazakhstan]] and [[Mongolia]], wolves are traditionally [[falconry|hunted using]] [[Golden eagles in human culture#Falconry|eagles]] and large falcons, though this practice is declining, as experienced falconers are becoming few in number. Shooting wolves from aircraft is highly effective, due to increased visibility and direct lines of fire.{{sfn|Graves|2007|pp=121–140}} Several types of dog, including the [[Borzoi]] and [[Taigan|Kyrgyz Tajgan]], have been specifically bred for wolf hunting.<ref name=Lescureaux2014/>

=== Wolves as pets and working animals ===

{{Main|Wolves as pets and working animals}}

Wolves and [[wolf-dog hybrid]]s are sometimes kept as [[exotic pet]]s, wolves do not show the same tractability as dogs in living alongside humans, being generally less responsive to human commands and more likely to act aggressively. Humans are more likely to be fatally mauled by a pet wolf or wolf-dog hybrid than by a dog.<ref name="Tucker1998"/>

== Notes ==

{{notelist}}

== References ==

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<ref name=Mech2003>{{cite book|last1=Mech|first1=L. David|last2=Adams|first2=L. G.|last3=Meier|first3=T. J.|last4=Burch|first4=J. W.|last5=Dale|first5=B. W.|title=The Wolves of Denali|publisher=University of Minnesota Press|year=2003|chapter=Ch.8-The Denali Wolf-Prey System|page=163|isbn=0-8166-2959-5|chapter-url={{Google books|plainurl=yes|id=-IZBwMrNWnMC|page=163}}}}</ref>

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<ref name=Werhahn2018>{{cite journal|doi=10.1016/j.gecco.2018.e00455|title=The unique genetic adaptation of the Himalayan wolf to high-altitudes and consequences for conservation|journal=Global Ecology and Conservation|volume=16|page=e00455|year=2018|last1=Werhahn|first1=Geraldine|last2=Senn|first2=Helen|last3=Ghazali|first3=Muhammad|last4=Karmacharya|first4=Dibesh|last5=Sherchan|first5=Adarsh Man|last6=Joshi|first6=Jyoti|last7=Kusi|first7=Naresh|last8=López-Bao|first8=José Vincente|last9=Rosen|first9=Tanya|last10=Kachel|first10=Shannon|last11=Sillero-Zubiri|first11=Claudio|last12=MacDonald|first12=David W.|doi-access=free|bibcode=2018GEcoC..1600455W |hdl=10651/50748|hdl-access=free}}</ref>
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}}

== Bibliography ==

*{{Cite book|last1=Busch|first1=R. H.|title=Wolf Almanac, New and Revised: A Celebration Of Wolves And Their World|publisher=[[Rowman & Littlefield]]|edition=3|year=2007|isbn=978-1-59921-069-8|url={{Google books|plainurl=yes|id=svbnCwAAQBAJ}}}}
*{{Cite book |last=Graves |first=Will |title=Wolves in Russia: Anxiety throughout the ages |publisher=Detselig Enterprises |year=2007 |isbn=978-1-55059-332-7 }}
*{{Cite book|last1=Heptner|first1=V. G.|last2=Naumov|first2=N. P.|url=https://archive.org/details/mammalsofsov211998gept|title=Mammals of the Soviet Union Vol. II Part 1a, Sirenia and Carnivora (Sea cows; Wolves and Bears)|publisher=Science Publishers, Inc. USA.|year=1998|isbn=978-1-886106-81-9}}
*{{Cite book |last=Lopez |first=Barry H. |title=Of Wolves and Men |publisher=J. M. Dent and Sons Limited |year=1978 |isbn=978-0-7432-4936-2 |url={{Google books|plainurl=yes|id=sz8gAQAAIAAJ}}}}
*{{Cite book |last=Marvin |first=Garry |title=Wolf |publisher=[[Reaktion Books]] Ltd |year=2012 |isbn=978-1-86189-879-1 |url={{Google books|plainurl=yes|id=7ib9WSIlvCsC}}}}
*{{Cite book |last=Mech |first=L. David |title=The Wolf: The Ecology and Behaviour of an Endangered Species |publisher=[[University of Minnesota Press]] |year=1981 |isbn=978-0-8166-1026-6 |url={{Google books|plainurl=yes|id=VqfwAAAAMAAJ}}}}
*{{Cite book|editor-last=Mech|editor-first=L. David|editor2-last=Boitani|editor2-first=Luigi|title=Wolves: Behaviour, Ecology and Conservation|publisher=[[University of Chicago Press]]|year=2003|isbn=978-0-226-51696-7|url={{Google books|plainurl=yes|id=zhwfmQEACAAJ}}}}
*{{Cite book|last1=Mech|first1=L. David|last2=Smith|first2=Douglas W.|last3=MacNulty|first3=Daniel R.|title=Wolves on the Hunt: The Behavior of Wolves Hunting Wild Prey|publisher=[[University of Chicago Press]]|year=2015|isbn=978-0-226-25514-9|url={{Google books|plainurl=yes|id=BJbICQAAQBAJ}}}}

== External links ==

{{Commons category|Canis lupus}}
{{Wikispecies|Canis lupus}}
{{Wikiquote|Wolf}}
*[https://www.wolf.org/ The International Wolf Center]
*[http://www.adfg.alaska.gov/static/species/livingwithwildlife/pdfs/wolf_safety_brochure.pdf Staying Safe in Wolf Country], [[Alaska Department of Fish and Game|ADFG]] (January 2009)
*Watch [https://www.nfb.ca/film/Death_of_a_Legend/ ''Death of a Legend''] and [http://www.nfb.ca/film/cry_of_the_wild/ ''Cry of the Wild''] {{Webarchive|url=https://web.archive.org/web/20090420215724/http://www.nfb.ca/film/cry_of_the_wild/ |date=April 20, 2009 }} by [[Bill Mason]]

{{Carnivora|Ca.}}
{{Canidae extinct nav|W.}}
{{Lupus}}
{{Mammal hybrids}}
{{Taxonbar|from=Q18498}}
{{Authority control}}

[[Category:Articles containing video clips]]
[[Category:Mammals of Asia]]
[[Category:Mammals of Europe]]
[[Category:Mammals of North America]]
[[Category:Extant Middle Pleistocene first appearances]]
[[Category:Fauna of the Holarctic realm]]
[[Category:Mammals described in 1758]]
[[Category:Pleistocene carnivorans]]
[[Category:Scavengers]]
[[Category:Taxa named by Carl Linnaeus]]
[[Category:Wolves| ]]
[[Category:Habitats Directive species]]
[[Category:Paraphyletic groups]]