Gray Wolf 33702 226155678 2008-07-17T01:59:13Z Seba5618 2155335 /* External links */ cleaning {{dablink|Wolf redirects here. For other uses, see [[Wolf (disambiguation)]], [[Gray Wolves (disambiguation)]], or [[Timber Wolf (comics)]].}} {{Taxobox | name = Gray Wolf | fossil_range = Late [[Pleistocene]] - Recent | status = LC | status_system = iucn3.1 | trend = stable | status_ref =<ref name="iucn">{{IUCN2006|assessors=Mech & Boitani|year=2004|id=3746|title=Canis lupus|downloaded=[[2006-05-05]]}} Database entry includes justification for why this species is of least concern.</ref> | image = Canis lupus signatus crop.jpg | image_caption = [[Iberian Wolf]] (''Canis lupus signatus'') | regnum = [[Animal]]ia | phylum = [[Chordate|Chordata]] | classis = [[Mammal]]ia | ordo = [[Carnivora]] | familia = [[Canidae]] | genus = ''[[Canis]]'' | species = '''''C. lupus''''' | binomial = ''Canis lupus'' | range_map= | range_map_caption=Range map. Green, present; red, former. | binomial_authority = [[Carolus Linnaeus|Linnaeus]], 1758 }} The '''gray wolf''' (''Canis lupus''), also known as the '''timber wolf''' or '''wolf''', is a [[mammal]] of the order [[Carnivora]]. The gray wolf is the largest wild member of the [[Canidae]] family and an [[ice age]] survivor originating during the [[Late Pleistocene]] around 300,000 years ago.<ref>Nowak, R. 1992. Wolves: The great travelers of evolution. International Wolf 2(4):3 - 7.</ref> [[DNA sequencing]] and [[genetic drift]] studies indicate that the gray wolf shares a common ancestry with the [[domestic dog]], (''Canis lupus familiaris'') and might be its ancestor.<ref>{{cite journal | author = Lindblad-Toh, K, ''et al.'' | title = Genome sequence, comparative analysis and haplotype structure of the domestic dog | journal = [[Nature (journal)|Nature]] | volume = 438 | pages = 803–819|year = 2005 | doi = 10.1038/nature04338}}</ref> A number of other gray wolf subspecies have been identified, though the actual number of subspecies is still open to discussion. Gray wolves are typically [[apex predators]] in the [[ecosystems]] they occupy. Gray wolves are highly adaptable and have thrived in temperate [[forests]], [[deserts]], [[mountains]], [[tundra]], [[taiga]], [[grasslands]] and [[urban areas]]. Though once abundant over much of [[North America]] and [[Eurasia]], the gray wolf inhabits a very small portion of its former range because of widespread destruction of its habitat, human encroachment of its habitat, and the resulting human-wolf encounters that sparked broad [[Local extinction|extirpation]]. Considered as a whole, however, the gray wolf is regarded as being of least concern for extinction according to the [[International Union for the Conservation of Nature and Natural Resources]]. Today, wolves are protected in some areas, hunted for sport in others, or may be subject to extermination as perceived threats to livestock and pets. In areas where human cultures and wolves are sympatric, wolves frequently feature in the folklore and mythology of those cultures, both positively and negatively. {{TOClimit|limit=3}} == Physiology == === Physical characteristics === Wolf weight and size can vary greatly worldwide, tending to increase proportionally with [[latitude]] as predicted by [[Bergmann's Rule]]. In general, height varies from 0.6 to .95 [[meter]]s (26–38 [[inch]]es) at the [[shoulder]] and weight ranges from 20 (44 lb.) to 62 (137 lb.) [[kilogram]]s, which together make the gray wolf the largest of all wild canids.<ref name="sinapu">{{cite web | url = http://www.yellowstone-bearman.com/wolves.html | title = Grey Wolves | publisher = Yellowstone-Bearman | accessdate = 2007-03-17 | month = November | year = 2002}}</ref> Although rarely encountered, extreme specimens of more than 77 kg (170 lb.) have been recorded in [[Alaska]], [[Canada]]<ref name="handbook">{{cite web | title = Persecution and Hunting | work = Endangered Species Handbook | publisher = Animal Welfare Institute | url = http://www.endangeredspecieshandbook.org/persecution_wolves2.php | accessdate = 2006-08-20}}</ref> and [[Russia]].<ref name="Graves">{{cite book | author = Graves, Will |url = http://www.wolvesinrussia.com/ | title = Wolves in Russia: Anxiety throughout the ages | year = 2007 | pages = pp.222 | id = ISBN 1550593323}}</ref> The heaviest recorded wild wolf in the [[New World]] was killed on 70 Mile River in east central Alaska on July 12, 1939 and weighed 79 kg (175 lb.)<ref name="SAM" />, while the heaviest recorded wild wolf in the [[Old World]] was killed after [[WWII]] in the kobelyakski Area of the Poltavskij Region in the [[Russian SFSR]] and weighed 86 kg (189 lb.).<ref name="Graves" /> The smallest wolves come from the [[Arabian Wolf]] subspecies, the females of which may weigh as little as 10 kg (22 lb) at maturity. Wolves are [[sexually dimorphic]], with females in any given wolf population typically weighing 20% less than males<ref name="WOLVES">{{cite book | author= L. David Mech & Luigi Boitani | title=Wolves: Behaviour, Ecology and Conservation | year=2001 | pages= p 448 | id= ISBN 0226516962 }}</ref>. They also have narrower muzzles and foreheads, slightly shorter, smoother furred legs and less massive shoulders<ref name="SAM" />. Wolves can measure anywhere from 1.3 to 2 meters (4.5–6.5 [[foot (unit of length)|feet]]) from nose to the tip of the [[tail (anatomy)|tail]], which itself accounts for approximately one quarter of overall body length.<ref>{{cite web | first= Angie | last= Hodgson | url= http://www.wcs.org/media/file/wcswp8.pdf | title= Wolf Restoration in the Adirondacks? | publisher= Wildlife Conservation Society | accessdate= 2006-08-21 | month= July | year= 1997 | format=PDF}}</ref> [[Image:WolfSkelLyd1.png|thumb|Wolf skeleton.]] Wolves are built for [[endurance|stamina]], possessing features ideal for long-distance travel. Their narrow chests and powerful backs and legs facilitate efficient [[Terrestrial locomotion in animals|locomotion]]. They are capable of covering several miles trotting at about a pace of 10 [[kilometer|km]]/h (6 [[miles per hour|mph]]), and have been known to reach speeds approaching 65 km/h (40 mph) during a chase.<ref>{{cite web | title = Gray Wolf Biologue | work = Midwest Region | publisher = U.S. Fish and Wildlife Service | url = http://www.fws.gov/midwest/wolf/biology/biologue.htm | accessdate = 2006-08-21}}</ref> One female wolf was recorded to have made 7 metre bounds when chasing prey.<ref name="Graves" /> Wolf [[paw]]s are able to tread easily on a wide variety of terrains, especially snow. There is a slight webbing between each toe, which allows them to move over snow more easily than comparatively hampered prey. Wolves are [[digitigrade]], which, with the relative largeness of their feet, helps them to distribute their weight well on snowy surfaces. The front paws are larger than the hind paws, and have a fifth digit, the [[dewclaw]], that is absent on hind paws.<ref name="WD">{{cite web | url = http://news.bbc.co.uk/1/hi/sci/tech/3602741.stm | title = "Claws reveal wolf survival threat" | work = Paul Rincon | publisher = BBC online | accessdate = 2007-05-11}}</ref> Bristled hairs and blunt claws enhance grip on slippery surfaces, and special [[blood vessel]]s keep paw pads from freezing.<ref name="AP">{{cite web | url = http://web.archive.org/web/20060524181949/http://animal.discovery.com/fansites/jeffcorwin/carnival/lilmammal/wolf.html | title = Gray Wolf | work = Corwin's Carnival of Creatures | publisher = Animal Planet | accessdate = 2006-05-24}}</ref> [[Scent gland]]s located between a wolf's toes leave trace chemical markers behind, helping the wolf to effectively navigate over large expanses while concurrently keeping others informed of its whereabouts.<ref name="AP" /> Unlike dogs and [[coyote]]s, wolves lack sweat glands on their paw pads. This trait is also present in Eastern Canadian Coyotes which have been shown to have recent wolf ancestry.<ref name="DOGS">{{cite book | author= Coppinger, Ray | title=Dogs: a Startling New Understanding of Canine Origin, Behavior and Evolution | year=2001 | pages= p352 | id=ISBN 0684855305 }}</ref> Wolves in [[Israel]] are unique due to the middle two toes of their paws being fused, a trait originally thought to be unique to the [[African Wild Dog]].<ref name="CLAW">{{cite book | author= Macdonald, David | title=The Velvet Claw | year=1992 | pages= pp.256 | id= ISBN 0563208449 }}</ref> [[Image:Wolfmoult.png|thumb|left|Wolves molt in late spring or early summer.]] Wolves have bulky [[coat (animal)|coats]] consisting of two layers. The first layer is made up of tough [[guard hair]]s that repel water and dirt. The second is a dense, water-resistant [[undercoat]] that insulates. The undercoat is shed in the form of large tufts of fur in late spring or early summer (with yearly variations). A wolf will often rub against objects such as rocks and branches to encourage the loose fur to fall out. The undercoat is usually gray regardless of the outer coat's appearance. Wolves have distinct winter and summer [[pelage]]s that alternate in spring and autumn. Females tend to keep their winter coats further into the spring than males. North American wolves typically have longer, silkier fur than their Eurasian counterparts.<ref name="LOU">{{cite book | author = Ellis, Shaun | title = Le Loup : Sauvage et Fascinant | year = 2006 | pages = pp.225 | id = ISBN 2749905389}}</ref> [[Fur]] coloration varies greatly, running from [[gray]] to gray-brown, all the way through the canine spectrum of white, red, brown, and black. These colors tend to mix in many populations to form predominantly blended individuals, though it is not uncommon for an individual or an entire population to be entirely one color (usually all black or all white). A multicolor coat characteristically lacks any clear pattern other than it tends to be lighter on the animal's underside. Fur color sometimes corresponds with a given wolf population's environment; for example, all-white wolves are much more common in areas with perennial snow cover. Aging wolves acquire a grayish tint in their coats. It is often thought that the coloration of the wolf's pelage serves as a functional form of camouflage. This may not be entirely correct, as some scientists have concluded that the blended colors have more to do with emphasizing certain gestures during interaction.<ref name="SAM">{{cite book | author = Lopez, Barry | title = Of wolves and men | year = 1978 | pages = pp.320 | id = ISBN 0743249364}}</ref> At birth, wolf pups tend to have darker fur and blue irises that will change to a yellow-gold or [[orange (colour)|orange]] color when the pups are between 8 and 16 weeks old.<ref name="IWCpups">{{cite web | url = http://www.wolf.org/wolves/learn/basic/biology/pupdevelopment.asp | title = Wolf Pup Development | work = Wolf Basics | publisher = International Wolf Center | accessdate = 2006-08-23 | month = November | year = 2004}}</ref> Though extremely unusual, it is possible for an adult wolf to retain its blue-colored irises.<ref>{{cite web | url= http://www.wolfspirits.org/wolves.htm | title= About Wolves | publisher= The Wolf Spirits | accessdate= 2006-08-21}}</ref> [[Image:Canis lupus pup closeup.jpg|thumb|right|Adolescent wolf with golden-yellow eyes.]] Wolves' long, powerful [[snout|muzzles]] help distinguish them from other canids, particularly [[coyote]]s and [[golden jackal]]s, which have more narrow, pointed muzzles. Wolves differ from domestic dogs in a more varied nature. Anatomically, wolves have smaller orbital angles than dogs ([[inequality|>]]53 [[degree (angle)|degrees]] for dogs compared with <45 degrees for wolves) and a comparatively larger brain capacity.<ref name="Evolution" /> Larger paw size, yellow eyes, longer legs, and bigger teeth further distinguish adult wolves from other canids, especially dogs. Also, precaudal glands at the base of the tail are present in wolves but not in dogs. Wolves and most larger dogs share identical [[dentition]]. The [[maxilla]] has six [[incisor]]s, two [[canine tooth|canines]], eight [[premolar]]s, and four [[molar (tooth)|molars]]. The [[mandible]] has six incisors, two canines, eight premolars, and six molars.<ref>{{cite web | title = The skull of Canis lupus | work = World of the Wolf | publisher = Natural Worlds | url = http://www.naturalworlds.org/wolf/moretopics/wolf_skull.htm|accessdate = 2005-08-21}}</ref> The fourth upper premolars and first lower molars constitute the [[carnassial]] teeth, which are essential tools for shearing flesh. The long canine teeth are also important, in that they hold and subdue the prey. Capable of delivering up to 10,000 [[kPa]] (1450 [[lbf/in²]]) of [[pressure]], a wolf's teeth are its main weapons as well as its primary tools.<ref name="SAM" /> The dentition of grey wolves is better suited to bone crushing than those of other modern canids, though it is not as specialised as that found in [[hyenas]].<ref name="Dire wolf">''Journal of Zoology'' Volume 267, Part 1, September 2005</ref> Wolf [[saliva]] has been shown to reduce bacterial infection in wounds and accelerate tissue regeneration.<ref name="Ellis" /> === Reproduction and life cycle === Usually, the instinct to reproduce drives young wolves away from their birth packs, leading them to seek out mates and [[territory (animal)|territories]] of their own. [[Biological dispersal|Dispersals]] occur at all times during the year, typically involving wolves that have reached [[sexual maturity]] prior to the previous [[breeding season]]. It takes two such dispersals from two separate packs for a new breeding pair to be formed, for dispersing wolves from the same maternal pack tend not to mate.<ref name="mating"/> Once two dispersing wolves meet and begin traveling together, they immediately begin the process of seeking out territory, preferably in time for the next mating season. The bond that forms between these wolves often lasts until one of them dies.<ref name="ADW">{{cite web | title = Canis lupus | work = Animal Diversity Web | publisher = University of Michigan Museum of Zoology | first = Tanya | last = Dewey | year = 2002 | url= http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_lupus.html | accessdate = 2005-08-18}}</ref> Generally, [[mating]] occurs between January and April — the higher the latitude, the later it occurs.<ref name="ADW" /> A pack usually produces a single litter unless the breeding male mates with one or more subordinate females. During the mating season, breeding wolves become very affectionate with one another in anticipation of the female's [[ovulation cycle]]. The pack tension rises as each mature wolf feels urged to mate. During this time, the breeding pair may be forced to prevent other wolves from mating with one another.<ref name="mating"> {{cite web | url = http://www.bio.davidson.edu/people/vecase/behavior/Spring2004/porter/Mating%20System.htm | title = Mating system | publisher = Department of Biology, Davidson College | accessdate = 2006-08-22}}</ref> Under normal circumstances, a pack can only support one [[litter (animal)|litter]] per year, so this dominance behavior is beneficial in the long run. When the breeding female goes into [[estrus]] (which occurs once per year and lasts 5–14 days),<ref>{{cite web | url= http://web.archive.org/web/20050505093101/http://www.dlia.org/atbi/species/animals/vertebrates/mammals/canidae/Canis_lupus.shtml | title= Gray Wolf | publisher= Discover Life in America|accessdate= 2005-05-05}}</ref> she and her mate will spend an extended time in seclusion. [[Pheromones]] in the female's [[urine]] and the swelling of her [[vulva]] make known to the male that the female is in heat. The female is unreceptive for the first few days of estrus, during which time she sheds the lining of her [[uterus]]; but when she begins ovulating again, the two wolves mate. The male wolf will mount the female firmly from behind. After achieving [[coitus]], the two form a copulatory tie once the male's [[bulbus glandis]]—an [[erectile tissue]] located near the base of the canine [[penis]]—swells and the female's [[vagina]]l muscles tighten. [[Ejaculation]] is induced by the thrusting of the male's [[pelvis]] and the undulation of the female's [[cervix]]. The two become physically inseparable for anywhere from 10 to 30 minutes, during which the male will ejaculate multiple times.<ref>{{cite web | url = http://www.mountainnature.com/Articles/WolvesCoyotes&Fox.htm | title = Wolves, Coyotes and Fox | publisher = MountainNature.com | accessdate = 2006-08-23}}</ref> After the initial ejaculation, the male may lift one of his legs over the female such that they are standing end-to-end; this is believed to be a defensive measure. The mating ritual is repeated many times throughout the female's brief ovulation period, which occurs once per year per female—unlike female dogs, whose estrus usually occurs twice per year. [[Image:Canis lupus resting in front of den.jpg|left|thumb|A wolf resting at the entrance to its den; also note how its coloration blends in with the environment.]] The [[gestation period]] lasts between 60 and 63 days. The pups, at a weight of 0.5 kg (1 lb), are born blind, deaf, and completely dependent on their mother.<ref name="ADW" /><ref name="sinapu" /> The average litter size is 5-6 pups, though litters of 10 can occur .<ref name="Graves" /> The pups reside in the [[den]] and stay there for two months. The den is usually on high ground near an open water source, and has an open chamber at the end of an [[underground living|underground]] or [[hillside]] [[tunnel]] that can be up to a few meters long.<ref name="AP" /> During this time, the pups will become more independent, and will eventually begin to explore the area immediately outside the den before gradually roaming up to a [[mile]] away from it at around 5 weeks of age. Wolf growth rate is slower than that of coyotes and [[dhole]]s.<ref name="DHOLE">{{cite book | author= Fox, Michael W. | title=The Whistling Hunters: Field Studies of the Asiatic Wild Dog (Cuon Alpinus) | year=1984 | pages= pp.150 | id=ISBN 0873958438 }}</ref> They begin eating [[Regurgitation (digestion)|regurgitated]] foods after 2 weeks of feeding on milk, which in wolves has less [[fat]] and more [[protein]] and [[arginine]] than dog milk.<ref name="WOLVES" /> By this time, their milk teeth have emerged — and are fully weaned by 10 weeks. During the first weeks of development, the mother usually stays with her litter alone, but eventually most members of the pack will contribute to the rearing of the pups in some way.<ref name="ADW" /> After two months, the restless pups will be moved to a [[rendezvous]] site, where they can stay safely while most of the adults go out to [[hunt]]. One or two adults stay behind to ensure the safety of the pups. After a few more weeks, the pups are permitted to join the adults if they are able, and will receive priority on anything killed, their low ranks notwithstanding. Letting the pups fight for eating privileges results in a secondary ranking being formed among them, and allows them to practice the dominance/submission rituals that will be essential to their future survival in pack life.<ref name="ADW" /> During hunts, the pups remain ardent observers until they reach about 8 months of age, by which time they are large enough to participate actively. Wolves typically reach [[sexual maturity]] after two or three years, at which point many of them will be compelled to leave their birth packs and seek out mates and territories of their own.<ref name="ADW" /><ref name="IWCagain">{{cite web | url = http://www.wolf.org/wolves/learn/basic/basic_main.asp | title = Gray Wolf Biology and Status | work = Wolf Basics | month = March | year = 2005|accessdate = 2006-08-23}}</ref> Wolves that reach maturity generally live 6 to 10 years in the [[wild]], although in [[captivity]] they can live to twice that age.<ref name="IWC">{{cite web | url= http://www.wolf.org/wolves/learn/basic/faqs/faq.asp | title= FAQ | work= Wolf Basics | publisher= International Wolf Center | first= Liz | last= Harper | month= November | year= 2002|accessdate= 2005-08-21}}</ref> High [[mortality rate]]s give them a low overall life expectancy. Pups die when food is scarce; they can also fall prey to predators such as [[bear]]s, or, less often, [[coyote]]s, or other wolves. The most significant causes of mortality for grown wolves are hunting and [[poaching]], [[car accident]]s, and wounds inflicted while hunting prey. Although adult wolves may occasionally be killed by other predators, rival wolf packs are often their most dangerous non-human enemy. A study on wolf mortality in Minnesota and the [[Denali National Park and Preserve]] concluded that 14–65% of wolf deaths were due to predation by other wolves.<ref>{{cite journal|title=Causes of wolf mortality in Croatia in the period 1986-2001|journal=Veterinarski Arhiv|date=|first=Đuro Huber|last=Huber|coauthors=Josip Kusak, Alojzije Frković, Goran Gužvica|volume=72|issue=3|pages=131–139|id= |url=http://www.environmental-studies.de/Wolf_mortality_Croatia.pdf|format=PDF|accessdate=2007-07-20}}</ref> ===Diseases=== Diseases recorded to be carried by wolves include [[rabies]], [[brucella]], [[deerfly fever]], [[listerosis]], [[foot and mouth disease]] and [[anthrax]]. Wolves are major hosts for rabies in Russia, [[Iran]], [[Afghanistan]], [[Iraq]] and [[India]]. Wolves in Russia have been recorded to carry over 50 different kinds of harmful parasites, including [[echinococcia]], [[cysticercocci]] and [[coenuri]]. Despite their habit of carrying harmful diseases, large wolf populations are not heavily regulated by epizootic outbreaks as with other social canids. This is largely due to the habit of infected wolves vacating their packs, thus preventing mass contagion.<ref name="Graves" /> == Behavior == === Social structure === Wolves function as social predators and hunt in [[pack (canine)|packs]] organized according to strict, rank-oriented [[social hierarchy|social hierarchies]].<ref name="ADW" /> Wolf packs tend not to be as compact or unified as those of [[African wild dog]]s and [[spotted hyena]]s<ref name="Hyena">{{cite book | author= Kruuk, Hans | title=The Spotted Hyena: A study of predation and social behaviour | year=1972 | pages= pp.335 | id= ISBN 0563208449 }}</ref>, though they are not as unstable as those of coyotes.<ref name="Encyclopedia">{{cite book | author = Macdonald, David | url = http://www.amazon.com/Encyclopedia-Mammals-David-W-Macdonald/dp/0871968711 | title = The Encyclopedia of Mammals: 1| year = 1984 | pages = pp.446 | id = ISBN 0-04-500028-x}}</ref> It was originally believed that this comparatively high level of social organization was related to hunting success, and while this still may be true to a certain extent, emerging theories suggest that the pack has less to do with hunting and more to do with [[reproduction|reproductive]] success. The pack is led by the two breeding individuals that sit atop the social hierarchy. The breeding pair has the greatest amount of social freedom compared to the rest of the pack. Although they are not "leaders" in the human sense of the term, they help to resolve any disputes within the pack, have the greatest amount of control over resources (such as food), and have exclusive rights to mating. While most breeding pairs are [[monogamy|monogamous]], there are exceptions.<ref>{{cite web | title = Wolf Family Life | publisher = Wolf Trust | url = http://www.wolftrust.org.uk/a-reproduction.html|accessdate = 2005-08-21}}</ref> An alpha animal may preferentially [[mating|mate]] with a lower-ranking animal, especially if the other alpha is closely related (a brother or sister, for example). The death of one breeding wolf does not affect the status of the other, who will quickly take another mate.<ref name="ADW" /> Usually, only the breeding pair is able to [[parenting|rear]] a litter of pups successfully. Other wolves in a pack may breed, but when resources are limited, time, devotion, and preference will be given to the alpha pair's litter. Therefore, non-alpha parents of other litters within a single pack may lack the means to raise their pups to [[maturity]] of their own accord. All wolves in a pack assist in raising wolf pups. Some mature individuals choosing not to disperse may stay in their original packs so as to reinforce it and help rear more pups. [[Image:Wolf pack in Yellowstone NP.jpg|thumb|left|A wolf pack in [[Yellowstone National Park]], with the breeding animals leading and the omega in the rear.]] After the breeding pair, there may also be a beta wolf whose rank is above that of the others save for the breeding pair. Betas typically assume a more prominent role in assisting with the upbringing of the breeding pair's litter, often serving as [[surrogate mother]]s or fathers while the breeding pair is away. Beta wolves are the most likely to challenge their superiors for the role of dominance, though some betas seem content with being second, and will sometimes even let lower ranking wolves [[leapfrog]] them for the position of breeding animal should circumstances necessitate such a happening, such as the death of the previous breeding animal. More ambitious beta wolves, however, will only wait so long before contending for breeding position unless they choose to disperse and create their own pack instead. Loss of rank can happen gradually or suddenly. An older wolf may simply choose to give way when a [[motivation|motivated]] challenger presents itself, yielding its position without bloodshed. On the other hand, the challenged individual may choose to fight back with varying degrees of intensity. While the majority of wolf [[aggression]] is ritualized and non-injurious, a high-stakes fight can easily result in [[injury]] for either or both parties. The loser of such a confrontation is frequently chased away from the pack or, rarely, may be killed as other aggressive wolves contribute to the insurgency. These types of confrontations are more common during the mating season. Deaths occasionally happen, with some dominant male wolves having been known to kill two to four wolves in his lifetime.<ref>{{cite web |url= http://www.isleroyalewolf.org/pe_wolf_kill.htm |title=Murder or Justified Canicide: East Pack kills alpha male of Chippewa Harbor Pack |publisher = The Wolves and Moose of Isle Royale | author=John Vucetich |date= [[January 31]], [[2006]] |accessdate=2007-08-10}}</ref> Rank order within a pack is established and maintained through a series of [[ritualization|ritualized]] [[combat|fights]] and posturing best described as "ritual bluffing". Wolves prefer ritualised displays of aggression to physical confrontations, meaning that high-ranking status is based more on [[Wiktionary:personality|personality]] or attitude than on size or [[physical strength]]. Rank, who holds it, and how it is enforced varies widely between packs and between individual animals. In large packs full of easy going wolves or in a group of [[juvenile (animal)|juvenile]] wolves, rank order may shift almost constantly, or even be circular (for instance, animal A dominates animal B, who dominates animal C, who dominates animal A). In a more typical pack, only one wolf will assume the role of the omega: the lowest-ranking member of a pack.<ref name="wolfpark" /> Omegas receive the most aggression from the rest of the pack, and may be subjected to different forms of truculence at any time—anything from constant dominance from other pack members to inimical, physical harassment. Submissive individuals are better suited for constant displays of active and passive submission than they are for living alone. Any form of camaraderie is preferable to solitude and, indeed, submissive wolves tend to choose low rank over potential starvation. Despite the aggression to which they are often subjected, omega wolves have also been observed to be among the most playful wolves in the pack, often enticing all of the members in a pack into chasing games and other forms of play. In general, omega wolves exist to help relieve pack tension. The size of the pack may change over time and is controlled by several factors, including habitat, personalities of individual wolves within a pack, and food supply. Packs can contain between 2 and 20 wolves, though 8 is a more typical size.<ref>{{cite web | title = Wolf Pack Size and Food Acquisition | work = Northern Prairie Wildlife Research Center | publisher = U.S. Geological Survey | url = http://www.npwrc.usgs.gov/resource/mammals/wpsize/main.htm | accessdate = 2005-08-21}}</ref> New packs are formed when a wolf leaves its birth pack, finds a mate, and claims a territory. Lone wolves searching for other individuals can travel very long distances seeking out suitable territories. Dispersing individuals must avoid the territories of other wolves because intruders on occupied territories are chased away or killed. Wolves acting unusually within the pack, such as [[epileptic]] pups or thrashing adults crippled by a trap or a gunshot, are usually killed by other members of their own pack.<ref name="SAM" /> === Body language === {{seealso|Dog communication}} [[Image:Grey Wolf 3.jpg|right|thumb|This facial expression is defensive and gives warning to other wolves to be cautious.]] [[Image:Grey Wolf 4.jpg|right|thumb|This facial expression shows fear.]] Wolves can [[Animal communication|communicate]] visually through a wide variety of expressions and moods ranging from subtle signals, such as a slight shift in weight, to more obvious ones, such as rolling on their backs to indicate complete submission.<ref>{{cite web | title = Communication | publisher = Wolfdancer Holding Company | url= http://www.wolfdancer.org/communication/ | accessdate = 2005-08-21}}</ref> * '''Dominance''' – A dominant wolf stands stiff legged and tall. The ears are erect and forward, and the hackles bristle slightly. Often the tail is held vertically and curled toward the back. This display asserts the wolf's rank to others in the pack. A dominant wolf may stare at a submissive one, pin it to the ground, "ride up" on its shoulders, or even stand on its hind legs. * '''Submission (active)''' – During active submission, the entire body is lowered, and the lips and ears are drawn back. Sometimes active submission is accompanied by muzzle licking, or the rapid thrusting out of the [[tongue]] and lowering of the hindquarters. The tail is placed down, or halfway or fully between the legs, and the muzzle often points up to the more dominant animal. The back may be partly arched as the submissive wolf humbles itself to its superior; a more arched back and more tucked tail indicate a greater level of submission. * '''Submission (passive)''' – Passive submission is more intense than active submission. The wolf rolls on its back and exposes its vulnerable [[throat]] and underside. The paws are drawn into the body. This posture is often accompanied by whimpering. * '''Anger''' – An angry wolf's ears are erect, and its fur bristles. The lips may curl up or pull back, and the incisors are displayed. The wolf may also arch its back, lash out, or [[snarl]]. * '''Fear''' – A frightened wolf attempts to make itself look small and less conspicuous; the ears flatten against the head, and the tail may be tucked between the legs, as with a submissive wolf. There may also be whimpering or barks of fear, and the wolf may arch its back. * '''Defensive''' – A defensive wolf flattens its ears against its head. * '''Aggression''' – An aggressive wolf snarls and its fur bristles. The wolf may crouch, ready to attack if necessary. * '''Suspicion''' – Pulling back of the ears shows a wolf is suspicious. The wolf also narrows its eyes. The tail of a wolf that senses danger points straight out, parallel to the ground. * '''Relaxation''' – A relaxed wolf's tail points straight down, and the wolf may rest [[sphinx]]-like or on its side. The wolf may also wag its tail. The further down the tail droops, the more relaxed the wolf is. * '''Tension''' – An aroused wolf's tail points straight out, and the wolf may crouch as if ready to spring. * '''Happiness''' – As dogs do, a wolf may wag its tail if in a joyful mood. The tongue may loll out of the mouth. * '''Hunting''' – A wolf that is hunting is tensed, and therefore the tail is horizontal and straight. * '''Playfulness''' – A playful wolf holds its tail high and wags it. The wolf may [[frolic]] and dance around, or bow by placing the front of its body down to the ground, while holding the rear high, sometimes wagged. This resembles the playful behavior of domestic dogs. === Howling and other vocalisations=== [[Image:Howlsnow.jpg|thumb|Howling adult wolf at the [[UK Wolf Conservation Trust]]]] Howling helps pack members keep in touch, allowing them to communicate effectively in thickly [[forest]]ed areas or over great distances. Howling also helps to call pack members to a specific location. Howling can also serve as a declaration of territory, as shown in a dominant wolf's tendency to respond to a human imitation of a "rival" wolf in an area the wolf considers its own. This behavior is stimulated when a pack has something to protect, such as a fresh kill. As a rule of thumb, large packs will more readily draw attention to themselves than will smaller packs. Adjacent packs may respond to each others' howls, which can mean trouble for the smaller of the two. Wolves therefore tend to howl with great care.<ref name="howl">{{cite web | title = What's in a Howl? | work = NOVA Online | publisher = PBS | url = http://www.pbs.org/wgbh/nova/wolves/howl.html | first = Fred H. | last = Harrington | month = November | year = 2000 | accessdate = 2005-08-21}}</ref> Wolves will also howl for communal reasons. Some scientists speculate that such group sessions strengthen the wolves' social bonds and camaraderie—similar to community [[singing]] among humans.<ref name="howl" /> During such choral sessions, wolves will howl at different [[timbre|tones]] and varying [[pitch (music)|pitches]], making it difficult to estimate the number of wolves involved. This confusion of numbers makes a listening rival pack wary of what action to take. For example, confrontation could be disastrous if the rival pack gravely underestimates the howling pack's numbers. A wolf's howl may be heard from up to ten miles away, depending on weather conditions. Observations of wolf packs suggest that howling occurs most often during the [[twilight]] hours, preceding the adults' departure to the hunt and following their return. Studies also show that wolves howl more frequently during the [[breeding season]] and subsequent rearing process. The pups themselves begin howling soon after emerging from their dens and can be provoked into howling sessions easily over the following two months. Such indiscriminate howling usually is intended for communication, and does not harm the wolf so early in its life.<ref name="howl" /> Howling becomes less indiscriminate as wolves learn to distinguish howling pack members from rival wolves. The Arabian and Iranian wolf subspecies are unusual as they are not known to howl.<ref name="SAM" /><ref>{{cite web | url= http://www.wolfsongalaska.org/wolves_in_iran.htm |title = The Iranian Wolf | publisher = Wolf Song of Alaska | accessdate = 2007-08-11}}</ref> [[Growling]], while teeth are bared, is the most visual warning wolves use. Wolf growls have a distinct, deep, [[Bass (musical term)|bass]]-like quality, and are often used to threaten rivals, though not necessarily to defend themselves. Wolves also growl at other wolves while being aggressively dominant. Wolves [[bark (dog)|bark]] when nervous or when they want to warn other wolves of danger but do so very discreetly and will not generally bark loudly or repeatedly as dogs do. Instead they use a low-key, breathy "whuf" sound to immediately get attention from other wolves. Wolves also "bark-howl" by adding a brief howl to the end of a bark. Wolves bark-howl for the same reasons they normally bark. Generally, pups bark and bark-howl much more frequently than adults, using these vocalizations to cry for attention, care, or food. A lesser known sound is the rally. Wolves will gather as a group and, amidst much tail-wagging and muzzle licking, emit a high-pitched wailing noise interspersed with something similar to (but not the same as) a bark. Rallying is often a display of submission to an alpha by the other wolves.<ref>[http://kiza.kcore.de/wolfpaper-archive/2001-02-07 Rally to Seneca + important information about Norwegian wolves], Wolfpaper Picture Archive (February 7, 2001)</ref> Wolves also whimper, usually when submitting to other wolves. Wolf pups whimper when they need a reassurance of security from their parents or other wolves. === Scent marking === [[Image:Duma roll.jpg|thumb|Wolves scent-roll to bring scents back to the pack.]] Wolves, like other canines, use [[scent marking]] to lay claim to anything—from territory to fresh kills.<ref name="wolfpark">{{cite web | title = Frequently Asked Questions About Wolves | publisher = Wolf Park | url = http://www.wolfpark.org/wolffaq.html|accessdate = 2006-08-22}}</ref> Alpha wolves scent mark the most often, with males doing so more than females. The most widely used scent marker is urine. Male and female alpha wolves urine-mark objects with a raised-leg stance (all other pack members squat) to enforce rank and territory. They also use marks to identify food caches and to claim kills on behalf of the pack. [[Defecation]] markers are used for the same purpose as urine marks, and serve as a more visual warning, as well.<ref name="wolfpark" /> Defecation markers are particularly useful for navigation, keeping the pack from traversing the same terrain too often and also allowing each wolf to be aware of the whereabouts of its pack members. Above all, though, scent marking is used to inform other wolves and packs that a certain territory is occupied, and that they should therefore tread cautiously. Wolves have [[scent gland]]s all over their bodies, including at the base of the tail, between toes, and in the eyes, [[genitalia]], and skin.<ref name="wolfpark" /> Pheromones secreted by these glands identify each individual wolf. A dominant wolf will "rub" its body against subordinate wolves to mark such wolves as being members of a particular pack. Wolves may also "paw" dirt to release pheromones instead of urine marking.<ref>{{cite web | url = http://fwie.fw.vt.edu/WWW/esis/lists/e059001.htm | publisher = Virginia Tech Conservation Management Institute | title = Species Wolf, Gray|date = [[1996-03-14]] | accessdate = 2006-08-23}}</ref> === Dietary habits === Wolves feed primarily on medium to large sized [[ungulate]]s, including [[Ovis|sheep]], [[caprinae|goats]], [[chamois]], [[pigs]], [[deer]], [[antelope]], [[caribou]], [[horse]]s, [[moose]], [[yak]], and [[bison]]. Other recorded large prey include [[marine mammals]] such as [[Pinniped|seals]]<ref name="Graves" /> and [[beached whale]]s.<ref name="AINU" /> Solitary wolves depend more on smaller animals, which they capture by pouncing and pinning with their front paws, though lone wolves have been recorded to bring down prey as large as bison unaided.<ref name="WOLVES" /> Some wolf packs in Alaska have been observed to feed on [[salmon]].<ref>[http://www.wildlifenews.alaska.gov/index.cfm?adfg=wildlife_news.view_article&articles_id=86 Alaska’s Salmon-Eating Wolves<!-- Bot generated title -->]</ref> They also prey on [[rodent]]s, [[game bird]]s and other small animals. A single wolf can eat up to 3.2-3.5 kg of food at a time, though they can eat as much as 13-15 kg when sufficiently hungry.<ref name="Graves" /> A wolf's yearly requirement is 1.5 tons of meat.<ref name="Graves" /> Wolves can go without sustenance for long periods of time, with one Russian record showing how one specimen survived for 17 days without food.<ref name="SAM" /> Research has shown that 2 weeks without food will not weaken a wolf's muscle activity.<ref name="Graves" /> After eating, wolves will drink large quantities of water to prevent [[uremic poisoning]].<ref name="SAM" /> A wolf's stomach can hold up to 7.5 litres of water.<ref name="Graves" /> [[Image:Canis lupus pack surrounding Bison.jpg|thumb|left|An [[American Bison]] standing its ground, thereby increasing its chance for survival.]] Wolf packs show little strategic cooperation in hunting unlike [[lion]]s, though wolf pairs have been shown to strategize when attacking large prey.<ref name="WOLVES" /> When hunting large prey, wolves typically attempt to conceal themselves as they approach the selected animal. Often, they will wait for the prey to pastourise, when it is distracted. Wolves generally do not engage in long chases, and will usually stop a pursuit after a chase of 10-180 metres, though there has been one documented case of a wolf chasing a moose for 36 km.<ref name="Graves" /> Wolves typically kill large prey by tearing at their [[haunch]]es and [[perinium]] areas, causing massive [[bleeding]].<ref name="Graves" /> Sometimes, the wolves will bite the throat, severing the [[windpipe]] or [[jugular]].<ref name="Sheep">{{cite web | url=http://www.aws.vcn.com/alaska.html | title = Wolf Predation on Sheep in Alaska | publisher = Abundant Wildlife Society of North America | accessdate = 2008-07-10}}</ref> A single bite can cause a wound up to 10-15 cm in length. A large deer in optimum health will succumb to three bites at the perinium area after a chase of 150 metres. Once the prey collapses, the wolves will tear open the abdominal cavity and commence feeding on the animal, sometimes whilst it is still alive.<ref name="Graves" /> On some occasions, wolves will not press an attack, and will wait for their prey to die from their wounds before commencing feeding.<ref name="Myths" /> Pack status is reinforced during feeding. The breeding pair usually eats first, starting with the [[heart]], [[liver]] and [[lungs]]. Wolves of intermediate rank will prevent low ranking animals from feeding until the dominant pair finishes eating.<ref name="Ellis" /> The [[stomach]] is eaten, though the contents are left untouched. The leg muscles are eaten next, with the hide and bones being the last to be eaten.<ref name="WOLVES" /> Though commonly portrayed as targeting solely sick or infirm animals,<ref name="IWCagain" /> there is little evidence that they limit themselves to such targets. Research from the former Soviet Union for example shows that in some cases, 93% of all killed prey may have no illnesses or infirmities. In the [[Nenets Autonomous Okrug]], wolves were shown to select pregnant female domestic caribou and calves rather than infirm specimens, with some reports showing that wolves bypassed emaciated, sickly animals altogether in favour of well fed ones.<ref name="Graves" /> However, most healthy, fit individuals will not run from wolves and will instead choose to stand their ground, thus increasing the possibility of injury to the attacking wolves. The wolves are more likely to yield when confronted by bold prey. Wolves are generally inefficient at killing large, assertive prey, with success rates as low as 20% which is due, in part, to the large size and defensive capabilities of their prey.<ref>{{cite web | url = http://www.sciencedaily.com/releases/2007/08/070830114825.htm | title = Wolves Find Happy Hunting Grounds In Yellowstone National Park | publisher = Science Daily | date = [[August 31]], [[2007]] | accessdate = 2007-09-17}}</ref> Wolves have on occasion been observed to engage in acts of [[surplus killing]]. An example of this was reported by a Conservation Officer for the Minnesota Department of Natural Resources, stating that during a spring snow storm, two wolves killed 21 deer, consuming only two.<ref name="Wolves&Hunting">{{cite web | url=http://www.aws.vcn.com/wolves_and_hunting.html | title = Wolves and Hunting | publisher = Abundant Wildlife Society of North America | accessdate = 2008-07-10}}</ref> Surplus killing in the wild peaks in winter months when heavy snow impedes the movements of large hooved prey.<ref name="Graves" /> Wolves will on occasion supplement their diet with vegetation, with some areas of the former Soviet Union reporting that wolves cause serious damage to [[watermelon]] plantations.<ref name="Graves" /> In certain localities in Eurasia where there is little natural prey, wolves will forage in [[garbage dump]]s. There are few cases of wolves in North America relying on garbage for food.<ref name="WOLVES" /> ===Interspecific predatory relationships=== Wolves typically dominate other canid species in areas where they are sympatric. In North America, wolves are generally intolerant of [[coyote]]s in their territory; two years after their re-introduction to the Yellowstone National Park, the wolves were responsible for a near 50% drop in coyote populations through both competition and predation.<ref name="Coyote" /> Wolves have been reported to dig coyote pups from their dens and kill them. Wolves typically do not consume the coyotes they kill. There are no records of coyotes killing wolves<ref name="WOLVES" />, though they have been known to gang up on wolves if they outnumber them.<ref name="Coyote">{{cite web |url=http://nationalzoo.si.edu/Publications/ZooGoer/1998/3/weavingwolfweb.cfm |title=Weaving A New Web: Wolves Change An Ecosystem |author=Jim Robbins |publisher=Smithsonian National Zoological Park |year=1998 |accessdate=2007-08-10}}</ref> Near identical interactions have been observed in [[Greece]] between wolves and [[golden jackal]]s.<ref name="GW">{{cite web | title = Conservation Action Plan for the golden jackal (Canis aureus) in Greece | work = | publisher = WWF Greece | url = http://www.lcie.org/Docs/Action%20Plans/Greece%20Golden%20Jackal%20Action%20Plan%202004.pdf | accessdate = 2007-07-31|format=PDF}}</ref> Wolves may kill [[foxes]], though not as frequently as they do with coyotes. [[Racoon dog]]s are also reportedly preyed upon.<ref name="WOLVES" /> [[Cougar]]s are encountered in North America. Wolves are usually hostile toward cougars and will kill kittens if given the opportunity. The wolf's relation to adult cougars is more complex. A pack often takes advantage of cougars, stealing kills and sometimes killing mature adults. Interactions between solitary wolves and cougars are rarer, but the two species have killed each other.<ref>{{cite web |url=http://www.ualberta.ca/~jzgurski/wild.html |title=Wolf Ecology: How wolves interact with other predators. |publisher=University of Alberta - Edmonton |accessdate=2007-08-10}}</ref> National Park Service cougar specialist Kerry Murphy stated that the cougar usually is at an advantage on a one to one basis, considering it can effectively use its claws, as well as its teeth, unlike the wolf which relies solely on its teeth. Yellowstone officials have reported that attacks between cougars and wolves are not uncommon. Multiple incidents of cougars taking wolves and vice versa have been recorded in Yellowstone National Park. However, researchers in Montana have found that wolves regularly kill cougars in the area, though they did not specify whether or not this was a pack situation.<ref>{{cite web |url= http://www.forwolves.org/ralph/wolves-deadcougar.htm |title=Park wolf pack kills mother cougar |publisher = Ralph Maughan's Wildlife Reports, The Wolf Recovery Foundation | date=[[April 12]], [[2003]] |author=Ralph Maughan |accessdate=2007-08-10}}</ref> [[Image:Canis lupus pack fighting Ursus arctos horribilis (illustration).jpg|thumb|Reconstruction of a wolf pack confronting a [[Grizzly bear]] by Adolph Murie (1944)]] [[Brown bear]]s are encountered in both Eurasia and North America. The majority of interactions between wolves and brown bears usually amount to nothing more than mutual avoidance. Serious confrontations depend on the circumstances of the interaction, though the most common factor is defence of food and young. Brown bears will use their superior size to intimidate wolves from their kills and when sufficiently hungry, will raid wolf dens. Brown bears usually dominate wolves on kills, though they rarely prevail against wolves defending den sites. Wolves in turn have been observed killing bear cubs, to the extent of even driving off the defending mother bears.<ref name="WOLVES" /> Deaths in wolf/bear skirmishes are considered very rare occurrences, the individual power of the bear and the collective strength of the wolf pack usually being sufficient deterrents to both sides.<ref>{{cite web |url=http://www.upress.umn.edu/mech/mech_chap1.html |title=The Wolves of Denali: Chapter 1 |author=L. David Mech, Layne G. Adams, Thomas J. Meier, John W. Burch, and Bruce W. Dale |publisher=University of Minnesota Press |accessdate=2007-08-10}}</ref> [[American black bear]]s occur solely in [[the Americas]], and interactions with wolves are much rarer than with brown bears, due to differences in habitat preferences. The majority of black bear encounters with wolves occur in the species' northern range, with no interactions being recorded in Mexico. Wolves have been recorded to kill black bears on numerous occasions without eating them. Unlike brown bears, black bears frequently lose against wolves in disputes over kills.<ref name="WOLVES" /> In areas where wolves and [[tiger]]s share ranges, such as the [[Russian Far East]], the two species typically display a great deal of dietary overlap, resulting in intense competition. Wolf and tiger interactions are well documented in [[Sikhote-Alin]], which until the beginning of the 20th century, held very few wolves. It is thought by certain experts that wolf numbers increased in the region after tigers were largely eliminated during the Russian colonization in the late 1800s and early 1900s. This is corroborated by native inhabitants of the region claiming that they had no memory of wolves inhabiting Sikohte-Alin until the 1930s, when tiger numbers decreased.<ref name="Alin" /> Tigers depress wolf numbers, either to the point of localized extinction or to such low numbers as to make them a functionally insignificant component of the ecosystem. Wolves appear capable of escaping competitive exclusion from tigers only when human persecution decreases the latters numbers.<ref name="Tigers&wolves">{{cite book | author = Matthiessen, Peter | title = Large Carnivores and the Conservation of Biodiversity: Biodiversity | year = 2005 | pages = pp.526 | id = ISBN 1559630809}}</ref> Today wolves are considered scarce in tiger inhabited areas, being found in scattered pockets, and usually as loners or small groups. First hand accounts on interactions between the two species indicate that tigers occasionally chase wolves from their kills, while wolves will scavenge from tiger kills. Tigers are not known to prey on wolves, though there are four records of tigers killing wolves without consuming them.<ref name="Alin">{{cite web | url = http://www.savethetigerfund.org/AM/Template.cfm?Section=Search1&template=/CM/ContentDisplay.cfm&ContentFileID=559 |title = Tigers and Wolves in the Russian Far East: Competitive Exclusion, Functional Redundancy, and Conservation Implications | publisher = savethetigerfund.org | accessdate = 2008-07-09}}</ref> Wolves may occasionally encounter [[striped hyena]]s in the Middle East, Central and Southern Asia, mostly in disputes over carcasses. Though hyenas usually dominate wolves on a one to one basis, wolf packs have been reported to displace lone hyenas from carcasses.<ref>{{cite web | url = http://www.hyaena.ge/striped.htm |title = Striped Hyaena | publisher = IUCN Species Survival Commission Hyaenidae Specialist Group | accessdate = 2007-08-11}}</ref> Wolf remains have been found in [[cave hyena]] den sites, though it is unknown if the wolves were killed or scavenged upon.<ref name="spelaea">{{cite web | url = http://64.233.183.104/search?q=cache:TH2j9N58LoQJ:nts1.cgu.cz/bulletin/contents/2006/vol81no4/237_diedrich.pdf+Crocuta+spelaea&hl=en&ct=clnk&cd=13| title = Prey and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823)in horizontal and vertical caves of the Bohemian Karst| work = CAJUSG. DIEDRICH & KARELŽÁK| publisher = | accessdate = 2008-01-20}}</ref> == Taxonomy == The gray wolf is a member of the [[genus]] ''[[Canis]]'', which comprises between 7 and 10 species. It is one of six species termed 'wolf', the others being the [[Red Wolf]] ''(Canis rufus)'', the [[Indian Wolf]] ''(Canis indica)'', the [[Himalayan Wolf]] ''(Canis himalayaensis)'', the [[Eastern Wolf]] ''(Canis lycaon)'' and the [[Ethiopian Wolf]] ''(Canis simensis)'', although concerning a couple of these there is still some uncertainty as to whether they should be considered subspecies of ''Canis lupus'' or species in their own right. Recent genetic research suggests that the Indian Wolf, originally considered only as a subpopulation of the [[Iranian Wolf]] (''Canis lupus pallipes''), represents a distinct species (''Canis indica''). Similar results were obtained for the [[Himalayan wolf]], which is traditionally placed into the [[Tibetan Wolf]] (''Canis lupus laniger'') <ref>R. K. Aggarwal, T. Kivisild, J. Ramadevi, L. Singh:''Mitochondrial DNA coding region sequences support the phylogenetic distinction of two Indian wolf species.'' Journal of Zoological Systematics and Evolutionary Research, Volume 45 Issue 2 Page 163-172, May 2007 [http://www.blackwell-synergy.com/doi/abs/10.1111/j.1439-0469.2006.00400.x online]</ref>. With respect to common names, [[American and British English spelling differences|spelling differences]] result in the alternative spelling ''grey wolf''. As the first-named and most widespread of species termed "wolf", gray wolves are often simply referred to as ''wolves''. It was one of the many species originally described by [[Carolus Linnaeus]] in his eighteenth-century work, ''[[Systema Naturae]]'', and it still bears its original classification, ''Canis lupus''.<ref>{{la icon}} {{cite book | last=Linnaeus | first=C | authorlink=Carolus Linnaeus | title=Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. | publisher=Holmiae. (Laurentii Salvii). | date=1758| pages=824 | url=http://dz1.gdz-cms.de/index.php?id=img&no_cache=1&IDDOC=265100}}</ref> The binomial name is derived from the [[Latin]] ''Canis'', meaning "dog", and ''lupus'', "wolf".<ref>{{cite book | last = Simpson | first = D.P. | title = Cassell's Latin Dictionary | publisher = Cassell Ltd. | date = 1979 | edition = 5 | location = London | pages = 883 | id = ISBN 0-304-52257-0}}</ref> [[Image:Canis lupus arabs head front.JPG|thumb|Desert dwelling grey wolf subspecies, such as this Arabian wolf, tend to be smaller than their more northern cousins.]] Classifying gray wolf subspecies can be challenging. Although scientists have proposed a host of [[subspecies]], wolf taxonomy at this level remains controversial.<ref>{{cite web | url = http://www.epa.gov/fedrgstr/EPA-SPECIES/2000/July/Day-13/e17621.htm | title = Endangered and Threatened Wildlife and Plants; Proposal To Reclassify and Remove the gray wolf From the List of Endangered and Threatened Wildlife in Portions of the Conterminous United States; Proposal To Establish Three Special Regulations for Threatened Gray Wolves End Hierarchical Links | work = Federal Register Environmental Documents | publisher = U.S. Environmental Protection Agency|date = [[2000-06-09]] | first = Stephen C. | last = Saunders | accessdate = 2006-08-23}}</ref> Indeed, only a single wolf species may exist. Taxonomic modification will likely continue for years to come. Current theories propose that the gray wolf first evolved in [[Eurasia]] during the early [[Pleistocene]]. The rate of changes observed in [[DNA]] sequence date the Asiatic lineage to about 800,000 years, as opposed to the American and European lineages which stretch back only 150,000.<ref>{{cite news | first= | last= | coauthors= | title=Indian wolves are world's oldest | date=[[2004-07-17]] | publisher=[[BBC News]] | url =http://news.bbc.co.uk/1/hi/world/south_asia/3804817.stm | work = | pages = | accessdate = 2007-07-19 | language = }}</ref> The gray wolf migrated into North America from the [[Old World]], probably via the [[Bering land bridge]], around 400,000 years ago. The gray wolf then coexisted with the [[Dire Wolf]] (''Canis dirus''). Although more heavily built and possessing a stronger bite, the dire wolf's dentition was less adept at crushing bones as the grey wolf was.<ref name="Dire wolf" /> The Dire Wolf ranged from southern Canada to South America until about 8,000 years ago when climate changes are thought to have caused it to become extinct. After that the gray wolf is thought to have become the prime canine predator in North America. === Subspecies === {{main|Subspecies of Canis lupus}} At one point, up to 50 gray wolf subspecies were recognized. Though no true consensus has been reached, this list can be condensed to 13–15 general extant subspecies. Modern classifications take into account the DNA, anatomy, distribution, and migration of various wolf colonies. [[As of 2005]], 37 subspecies are currently described, including the [[dingo]] and the domestic dog.<ref>{{MSW3 Wozencraft | pages = | id = }}</ref> === Relation to the dog === {{Main|Origin of the domestic dog}} [[Image:DogWolf.jpg||thumb|Comparative drawing of dog and wolf anatomy by [[Ernest Thompson Seton]]. Note the proportionately larger head of the wolf]] Much debate has centered on the relationship between the wolf and the domestic dog, though most authorities see the wolf as the dog's direct [[ancestor]]. Because the canids have evolved recently and different canids interbreed readily, untangling the relationships has been difficult. However, [[molecular systematics]] now indicate very strongly that domestic dogs and wolves are closely related, and the domestic dog is now normally classified as a [[subspecies]] of the wolf: ''Canis lupus familiaris''. All skeletal dog remains found from the upper Paleolithic and Mesolithic periods are from relatively small specimens, therefore pointing to either the Arabian or Iranian wolf as the most likely progenitor.<ref name="Domestic" /> North American domestic dogs are believed to have originated from Old World wolves. No known dog breed is derived from wolves indigenous to North America. The first people to colonize North America 12,000 to 14,000 years ago brought their dogs with them from Asia, and apparently did not separately domesticate the wolves they found in the New World.<ref>{{cite web |url=http://nationalzoo.si.edu/ConservationAndScience/SpotlightOnScience/fleischer2003108.cfm |title= Spotlight on Zoo Science: Hiding In Plain Sight |accessdate=2007-10-07 |author=Susan Lumpkin |date= [[2003-10-08]] |work= Conservation and Science |publisher=Smithsonian National Zoological Park}}</ref> Compared to equally sized wolves, dogs tend to have 20% smaller skulls and 10% smaller brains, as well as proportionately smaller teeth than other canid species.<ref name="DOGS">{{cite book | author= Coppinger, Ray | title=Dogs: a Startling New Understanding of Canine Origin, Behavior and Evolution | year=2001 | pages= p352 | id=0684855305 }}</ref> The premolars and molars of a dog are much more crowded and compacted than those of a wolf. Dog's teeth also have less complex cusp patterns, and their [[tympanic bulla]] is much smaller than in wolves.<ref name="Domestic">{{cite book | author = Clutton-Brock, Juliet| title = A Natural History of Domesticated Mammals | year = 1987 | pages = pp.208 | id = ISBN 0521346975}}</ref> Dogs require fewer calories to function than wolves. The dog's diet of human refuse in antiquity made the large brains and jaw muscles needed for hunting unnecessary. It is thought by certain experts that the dog's limp ears are a result of [[atrophy]] of the jaw muscles.<ref name="DOGS" /> The paws of a dog are half the size of those of a wolf, and their tails tend to curl upwards, another trait not found in wolves.<ref name="SAM" /> Dogs are not [[monogamous]], and breeding in feral packs is not restricted to a dominant breeding pair. Male dogs differ from male wolves by the fact that they play no role in raising their puppies, and do not kill the young of other females to increase their own reproductive success.<ref name="Evolution">{{cite book | author= Serpell, James| title=The Domestic Dog; its evolution, behaviour and interactions with people | year=1995 | pages= p267 | id=0-521-42537-9 }}</ref> Dogs differ also from wolves by the fact that they do not regurgitate food for their young, nor the young of other dogs in the same territory.<ref name="DOGS" /> === Interspecific hybridization === {{Main|Canid hybrid}} [[Image:Wolf-malamute hybrid.jpg|thumb|A [[wolf-dog hybrid]] with [[malamute]] ancestry]] Wolves can interbreed with domestic dogs and produce fertile offspring. [[Wolf-dog hybrid]]s are generally said to be naturally healthy animals, and are affected by less inherited diseases than most breeds of dog. Wolfdogs are usually healthier than either parent due to [[heterosis]].<ref name="Controversy">{{cite web|date=2000|title=The Wolf-Dog Hybrid: An Overview of a Controversial Animal|publisher= Animal Welfare Information Center Newsletter|url=http://www.nal.usda.gov/awic/newsletters/v5n4/5n4wille.htm|accessdate=2008-05-17}}</ref> According to the [[National Wolfdog Alliance]], 40 U.S. states effectively forbid the ownership, breeding and importation of wolfdogs, while others impose some form of regulation upon ownership.<ref>[http://www.wolfdogalliance.org/legislation/statelaws.html NWA]</ref> Most European nations, as well as many U.S. counties and municipalities, also either outlaw the animal entirely or put restrictions on ownership.<ref>[http://www.wolftrust.org.uk/wolfdogs.html Wolf Trust]</ref><ref>[http://www.wolfdogalliance.org/legislation/statelaws.html National Wolf Dog Alliance]</ref>. Although wolves in the wild will usually kill dogs, matings of dogs and wild wolves has been confirmed in some populations through genetic testing. As the survival of most Continental wolf packs is severely threatened, scientists fear that the creation of wolf-dog hybrid populations in the wild is a threat to the continued existence of some isolated wolf populations. Hybridization in the wild usually occurs near human habitations where wolf density is low and dogs are common. However, extensive wolf-dog hybridization is not supported by morphological evidence, and analyses of [[mtDNA]] sequences have revealed that such matings are rare.<ref name="WOLVES" /> In some cases, the presence of [[dewclaw]]s is considered a useful, but not absolute indicator of dog gene contamination in wild wolves. Dewclaws are the vestigial fifth toes of the hind legs common in domestic dogs but thought absent from pure wolves, which only have four hind toes.<ref name="WD" /> Observations on wild wolf hybrids in the former Soviet Union indicate that wolf hybrids in a wild state may form larger packs than pure wolves, and have greater endurance when chasing prey.<ref name="Graves" /> Wolves and coyotes can interbreed and produce fertile offspring, a fact which calls into question their status as two separate species.<ref>[http://www.newscientist.com/channel/life/mg19726441.900-the-decline-fall-and-return-of-the-red-wolf.html The decline, fall and return of the red wolf - life - 23 February 2008 - New Scientist<!-- Bot generated title -->]</ref> The offspring, known as a [[coywolf]], is generally intermediate in size to both parents, being larger than a pure coyote, but smaller than a pure wolf. A study showed that of 100 coyotes collected in [[Maine]], 22 had half or more wolf ancestry, and one was 89 percent wolf. A theory has been proposed that the large eastern coyotes in Canada are actually hybrids of the smaller western coyotes and wolves that met and mated decades ago as the coyotes moved toward New England from their earlier western ranges.<ref name="EC">{{cite web | title = Eastern Coyotes Are Becoming Coywolves | work = David Zimmerman | publisher = Caledonian record | url = http://www.caledonianrecord.com/pages/local_news/story/fef373e9d | accessdate = 2007-08-17}}</ref> ==Current status== ===Europe=== [[Greenland]] has a population of 50-100 wolves which are afforded protection in approximately 90% of their range, though no compensation is paid for livestock damages.<ref name="WOLVES" /> [[Portugal]] has a stable wolf population of 200-300 which is afforded full protection. Compensation is paid for livestock damages.<ref name="WOLVES" /> [[Spain]]'s wolf population is estimated at 2000 and growing. Wolves are considered a game species, though they are protected in the southern regions of the country. Compensation is paid for livestock damages, though this varies according to regional laws.<ref name="WOLVES" /> In [[Italy]], wolves are a protected species, with current estimates indicating that there are 500-800 wolves living in the wild. The largest concentrations of wolves occur in the [[alps]] and the [[Italian national park]]s of [[Parco nazionale del Pollino|Pollino]], [[Parco Nazionale d'Abruzzo, Lazio e Molise|Abruzzo, Lazio e Molise]] and [[Appennino Tosco-Emiliano]]. Isolated individuals have been sighted in the vicinity of human populated areas such as [[Tuscany]], [[Rome]], [[Bologna]], [[Parma]] and [[Tarquinia]]. Currently, their populations are said to have been increasing at a rate of 6% a year since the 1970s, though 15% of the total Italian wolf population is reported to succumb annually to illegal poaching and road accidents.<ref name="arca">{{it icon}}{{cite book | author= Luigi Boitani & Paolo Ciucci | title="Il Lupo: Operazione San Francesco" from "Salvati dall' Arca" | year=2006 | pages= pp.663 | id=ISBN 9788883723803 }}</ref> Compensation is paid by regional governments for livestock damages.<ref name="WOLVES" /> Wolves migrated from Italy to [[France]] as recently as [[1992]], and the current French wolf population is said to be composed of 40-50 individuals and growing.<ref name="WOLVES" /> Under the [[Convention on the Conservation of European Wildlife and Natural Habitats|Berne Convention]], wolves are listed as an endangered species and killing them is illegal. Official culls are permitted to protect farm animals so long as there is no threat to the national population as a whole.<ref>[http://www.wolfsongalaska.org/gray_wolf_europe_cull.htm Wolf Song of Alaska: France's Bardot Demands EU Action on Wolf Cull<!-- Bot generated title -->]</ref> Compensation is paid for livestock damages.<ref name="WOLVES" /> Wolves were first spotted in [[Germany]] in 1998, and are thought to have migrated from western [[Poland]].<ref name="GER(b)">{{cite web | url=http://www.spiegel.de/international/spiegel/0,1518,467205,00.html | title = The Return of Wolves to Germany | work = | publisher = Spiegel online | accessdate = 2007-02-07}}</ref> Currently, there are around 35 wolves in 4 packs now roaming the heaths of the eastern German region of [[Lusatia]], and they are now still expanding their range to the west and north.<ref Name="GER(a)">{{cite web | url=http://www.wolfsregion-lausitz.de/index.php?article_id=12&clang | title = Aktueller Bestand in der Lausitzer Wolfsregion | work = | publisher = Wolfsregion Lausitz online | accessdate = 2007-11-02}}</ref> Under German law, wolves are a protected species, though there are no livestock damage compensation programmes.<ref name="WOLVES" /> The number of wolves in [[Switzerland]] is uncertain, having been guessed at 1-2 individuals. Wolves are afforded protection, and livestock damage compensation is paid by [[Cantons of Switzerland|Cantons]].<ref name="WOLVES" /> [[Norway]] currently has a stable, protected population of 15-20 wolves which is increasing. Compensation is paid for livestock damages.<ref name="WOLVES" /> [[Sweden]] has a protected population of 70-80 wolves and growing, and compensation is paid for livestock damages.<ref name="WOLVES" /> [[Finland]] has a stable population of 100 wolves which is increasing. Wolves are legally hunted only in areas with high reindeer densities. Compensation for livestock losses are paid by the state and insurance companies.<ref name="WOLVES" /> [[Poland]] has an increasing population of 700-800 wolves which are afforded legal protection except in the [[Bieszczady Mountains]]. Compensation for livestock losses is not paid.<ref name="WOLVES" /> [[Estonia]] is thought to have a wolf population of 500, though it is decreasing as the species is outlawed, and no livestock compensation is in effect, as the insurance is considered too expensive.<ref name="WOLVES" /> [[Lithuania]] has over 600 wolves which are increasing in number. The species is not protected, and only insured livestock receives compensation.<ref name="WOLVES" /> [[Latvia]] has an unprotected, yet stable population of 900 wolves. No compensation is paid for livestock damages.<ref name="WOLVES" /> [[Belarus]] is home to an increasing population of 2,000-2,500 wolves.<ref name="WOLVES" /> With the exception of specimens in nature reserves, wolves in Belarus are largely unprotected. They are designated a game species, and bounties ranging between €60 and €70 are paid to hunters for each wolf killed. This is a considerable sum in a country where the average monthly wage is €230.<ref>http://www.wolf.org/wolves/news/iwmag/2006/fall/wow_belarus.pdf</ref> No compensation is paid for livestock losses.<ref name="WOLVES" /> [[Ukraine]] has an unprotected, yet stable population of 2,000 wolves. No compensation is paid for livestock losses.<ref name="WOLVES" /> The [[Czech Republic]] has a stable and protected population of 20 wolves, though there are no livestock damage compensation programmes.<ref name="WOLVES" /> [[Slovakia]] has a stable population of 350-400 wolves which is protected, though with some exceptions. No compensation is paid for livestock losses.<ref name="WOLVES" /> [[Croatia]] has a population of 100-150 wolves and increasing. As of May 1995, they are a protected species, and compensation is paid for livestock losses.<ref name="WOLVES" /> The [[Federation of Bosnia and Herzegovina]] is thought to have a population of 400 wolves, though they are decreasing in number and are afforded no legal protection. Compensation for livestock losses is not paid.<ref name="WOLVES" /> [[Serbia and Montenegro]] has a stable population of 500 wolves, though it is unknown if they are afforded any protection and no compensation is paid for livestock damages.<ref name="WOLVES" /> [[Hungary]] has a stable population of 50 wolves which are protected, though with some exceptions. No compensation is paid for livestock damages.<ref name="WOLVES" /> [[Romania]] has a increasing population of 2,500 wolves which are granted legal protection, though no compensation is paid for livestock damages.<ref name="WOLVES" /> [[Bulgaria]] has a stable population of 800-1,000 wolves which are granted no legal protection.<ref name="WOLVES" /> Wolves are considered a nuisance and have an active bounty on them.<ref name="UKWCT">{{cite web | url = http://ukwolf.org/uk-wolf/71 | title = Bulgaria | work = | publisher = | accessdate = 2007-09-10}}</ref> No compensation is paid for livestock damages.<ref name="WOLVES" /> [[Greece]] has a stable population of 200-300 wolves which are legally protected. Compensation is paid for livestock losses, with over 80% of it from insurance.<ref name="WOLVES" /> The [[Republic of Macedonia]] has an increasing, yet unprotected population of 1,000 wolves, with no livestock compensation programmes.<ref name="WOLVES" /> [[Albania]] has a protected population of 250 wolves which are increasing in number, though no compensation is paid for livestock losses.<ref name="WOLVES" /> [[Turkey]] has an unknown number of wolves thought to be as high as 1,000. It is not known if they are increasing or decreasing, and no legal protection nor livestock damage compensation is granted.<ref name="WOLVES" /> Although wolves in [[Russia]] have no legal protection, they number 25,000-30,000, and are increasing in number.<ref name="WOLVES" /> No livestock damage compensation is paid.<ref name="WOLVES" /> ===Asia=== [[Syria]] has an unprotected, unknown number of wolves, thought to be roughly numbering 200. No livestock damage compensation is paid.<ref name="WOLVES" /> [[Lebanon]] has a population of 50 wolves which are afforded no legal protection, nor is livestock damage compensation is paid.<ref name="WOLVES" /> [[Israel]] has a stable population of 150 unprotected wolves. No livestock damage compensation is paid.<ref name="WOLVES" /> [[Jordan]] has an unprotected, unknown number of wolves, thought to be roughly numbering 200. No livestock damage compensation is paid.<ref name="WOLVES" /> [[Saudi Arabia]] has a stable population of 300-600 wolves which are given no legal protection. No livestock damage compensation is paid.<ref name="WOLVES" /> [[India]] has a decreasing population of roughly 1,000 wolves which are legally protected. No livestock damage compensation is paid.<ref name="WOLVES" /> The [[People's Republic of China]] considers wolves a "catastrophe" and claims that they live in only twenty percent of their former habitat in the northern regions of the country.<ref>[http://www.care2.com/c2c/groups/disc.html?gpp=2774&pst=172295 Wolf Subspecies - Wolflovers Group - Care2.com<!-- Bot generated title -->]</ref> Wolves in China appear to be decreasing in all their ranges. Currently, [[Cheiludjiang]] has roughly 500 wolves, [[Xinjiang]] has 10,000 and [[Tibet]] has 2,000.<ref name="WOLVES" /> In [[2006]], the Chinese government began plans to auction licenses to foreigners to hunt wild animals, including wolves.<ref>[http://news.bbc.co.uk/2/hi/asia-pacific/4775335.stm BBC NEWS | Asia-Pacific | China to promote wild animal hunt<!-- Bot generated title -->]</ref> No livestock damage compensation is paid.<ref name="WOLVES" /> [[Mongolia]] has a stable population of 10,000-20-000 wolves which are given no legal protection, nor is livestock damage compensation is paid.<ref name="WOLVES" /> [[Kazakhstan]] has a stable population of about 30,000 wolves.<ref name="WOLVES" /> About 2,000 are killed yearly for a $40 bounty, though the animal’s numbers have risen sharply.<ref name="KAZ">{{cite web | url = http://www.nwf.org/nationalwildlife/article.cfm?issueID=115&articleID=1473 | title = Is Kazakhstan Home to the World’s Largest Wolf Population? | work = Christopher Pala | publisher = National Wildlife Federation | accessdate = 2007-09-28}}</ref> No livestock damage compensation is paid.<ref name="WOLVES" /> [[Turkmenistan]] has a stable population of 1,000 wolves which are unprotected. No livestock damage compensation is paid.<ref name="WOLVES" /> [[Uzbekistan]] has a stable population of 2,000 wolves which are unprotected. No livestock damage compensation is paid.<ref name="WOLVES" /> [[Kirgizstan]] has a stable population of 4,000 wolves which are unprotected. No livestock damage compensation is paid.<ref name="WOLVES" /> [[Tadjikistan]] has a stable population of 3,000 wolves which are unprotected. No livestock damage compensation is paid.<ref name="WOLVES" /> There are currently no recent or reliable estimates on wolf populations in [[Iraq]], [[Iran]], [[Afghanistan]], [[Nepal]] and [[Bhutan]].<ref name="WOLVES" /> ===North America=== The [[United States]] as a whole has up to 9,000 wolves which are increasing in number in all their ranges. Wolf recovery has been so successful that the [[United States Fish & Wildlife Service]] removed the western grey wolf from the federal endangered species list on March 28, 2008.<ref name ="FWSWOLF">{{cite web | url = http://www.fws.gov/mountain-prairie/species/mammals/wolf/ | title = U.S. Fish & Wildlife Service Gray Wolf webpage.| work = | publisher = U.S. Fish & Wildlife Service | accessdate = 2008-04-23}}</ref><ref name="WOLVES" /> Due to the controversy over wolf shootings, a coalition of environmental groups is planning to sue the federal government to put the gray wolf back on the [[United States Fish and Wildlife Service list of endangered species|Endangered Species list]].<ref name = "NYTWOLF">{{cite web | url = http://www.nytimes.com/2008/04/13/us/13wolves.html?_r=1&ref=environment&oref=slogin | title = New York Times article on gray wolf controversy, 4/13/08 | work = | publisher = New York Times | accessdate = 2008-04-23}}</ref> [[Alaska]] has a stable population of 6,000-7,000 wolves which are legally hunted from August to April as a big game species.<ref name="WOLVES" /><ref name="Wolf Status World">{{cite web | url = http://www.dnr.state.mn.us/mammals/wolves/mgmt.html | title = Wolf management| work = | publisher = Minnesota Department of Natural Resources | accessdate = 2008-01-13}}</ref> Aerial hunting of wolves and other predators is used as a method to boost moose populations for hunters in Alaska.<ref name="nyt_alaska">{{cite web | url = http://query.nytimes.com/gst/fullpage.html?res=9801E7D7113EF937A25750C0A9629C8B63 | title = Wolf 'Control' in Alaska| work = | publisher = The New York Times | accessdate = 2008-01-13}}</ref> This practice is controversial. Biologists have cited possibly flawed scientific logic in opposing aerial hunting, but the citizens of Alaska have twice voted against aerial hunting.<ref name="nyt_alaska" /> [[Minnesota]] has a population of 2,500 wolves which are legally protected, though they are occasionally culled for depredation control. Minnesota is currently the only US state to have a livestock damage compensation programme.<ref name="WOLVES" /> Minnesota has been granted complete control over its wolf population, and its wolf management plan establishes a minimum population of 1600 wolves.<ref name="MNDNR">{{cite web | url = http://www.ualberta.ca/~jzgurski/population.html | title = Wolf populations around the world and their status.| work = | publisher = | accessdate = 2008-01-13}}</ref> [[Montana]] has population of 70 wolves which are legally protected.<ref name="WOLVES" /> [[Idaho]] has a population of 185 wolves which though protected, is considered merely experimental and nonessential.<ref name="WOLVES" /> [[Wyoming]] has 165 wolves, which like in Idaho, are considered merely experimental.<ref name="WOLVES" /> [[Michigan]] has 200 wolves which are legally protected.<ref name="WOLVES" /> [[Wisconsin]], like Michigan, has a population of 200 legally protected wolves.<ref name="WOLVES" /> [[Canada]] has over 52,000-60,000 wolves which are legally considered a big game species, though they are afforded protection in 3% of Canada's territory. The [[Northwest Territories]], [[Nunavut]] and [[Yukon]] have 5,000 wolves each, [[British Columbia]] has 8000 wolves, [[Alberta]] 4,200, [[Saskatchewan]] 4,300, [[Manitoba]] 4,000-6,000, [[Ontario]] 9,000, [[Quebec]] 5,000 and [[Labrador]] 2,000. Canada currently has no livestock damage compensation programmes.<ref name="WOLVES" /> [[Mexico]] has been wolf-free since the 1970s when the U.S. and Mexican governments cooperated to capture all remaining wild Mexican wolves and initiate a captive-breeding program in an attempt to save the local subspecies.<ref name = Mexwolfstats /> The Mexican Wolf was reintroduced into the [[Apache-Sitgreaves National Forest]] in Arizona in 1998 as part of a captive breeding program.<ref name = "Wolf Status World" /> There are now at least 59 wild Mexican wolves in the American southwest.<ref name="Wolf Status World" /><ref name = Mexwolfstats>{{cite web | url = http://www.fws.gov/southwest/es/mexicanwolf/pdf/Table_1_Mx_wolf_population_count_03082007.pdf | title = Mexican Wolf Blue Range Reintroduction Project Statistics| work = | publisher = USFWS | accessdate = 2008-01-13|format=PDF}}</ref> ==Relationships with humans== Humans historically have had a complex relationship with wolves. In many parts of the world, wolves were respected and revered, while in others they were feared and held in distaste. The latter viewpoint was notably accentuated in European [[folklore]] beginning in the [[Christian]] era, though wolves did feature as [[heraldry|heraldic]] animals on the [[Coats of arms|Arms]] and [[family crest|crest]]s of numerous noble families. Many languages have names meaning "wolf", examples including: [[Scandinavian languages|Scandinavian]] ''Ulf'', [[Albanian language|Albanian]] "Ujk", [[Hebrew]] ''Ze'ev'', [[Hungarian language|Hungarian]] ''Farkas'', [[Serbian language|Serbian]] ''Vuk'', [[Ukrainian language|Ukrainian]] ''Vovk'', [[Romanian language|Romanian]] ''Lupu, Lupescu/Lupulescu'' and [[Bulgarian language|Bulgarian]] ''Vǎlko''. Wolves also figure prominently in [[proverb]]s. Many [[Chinese proverb]]s use wolves as a description towards any ill-willed person with a hidden agenda like ''Wolf hearted'' (狼子野心) which could also connote to the impossibility of taming bad people, while ''Wolf heart; dog lungs'' (狼心狗肺) refers to an ungrateful person who later betrays someone who previously helped them. The [[Kazakh language]] has up to 20 proverbs referring to wolves, while the [[Russian language]] has 253.<ref name="Graves"/> ===In folklore and mythology=== {{Main|Wolves in folklore, religion and mythology}} [[Image:Romolo e remo.jpg|thumb|right|[[Romulus and Remus]] nursed by the [[Capitoline Wolf|She-wolf]] by [[Peter Paul Rubens]] ([[Rome]], [[Capitoline Museums]])]] In [[Altaic mythology]] of the [[Turkic peoples|Turkic]] and [[Mongolian peoples]], the wolf is a revered animal. The shamanic Turkic peoples even believed they were descendants of wolves in Turkic legends. The legend of [[Asena]] is an old Turkic myth that tells of how the Turkic people were created. In Northern China a small Turkic village was raided by Chinese soldiers, but one small baby was left behind. An old she-wolf with a sky-blue mane named Asena found the baby and nursed him, then the she-wolf gave birth to half wolf, half human cubs therefore the Turkic people were born. Also in Turkic mythology it is believed that a gray wolf showed the Turks the way out of their legendary homeland ''Ergenekon'', which allowed them to spread and conquer their neighbours.<ref>[http://www.byegm.gov.tr/yayinlarimiz/kitaplar/turkiye2003/content/english/434-435.htm Cultural Life – Literature] Turkey Interactive CD-ROM. Retrieved on [[2007-08-11]].</ref><ref> T.C. Kultur Bakanligi. [http://www.discoverturkey.com/english/kultursanat/h-nevruz.html Nevruz Celebrations in Turkey and Central Asia.] Ministry of Culture, Republic of Turkey. Retrieved on [[2007-08-11]],</ref> In modern Turkey this myth inspired extreme-right nationalist groups known as "[[Grey Wolves]]". The genesis story of the Turks and Mongols is paralleled in the Roman myth of [[Romulus and Remus]], the traditional founders of [[Rome]]. The twin babies were ordered to be killed by their great uncle [[Amulius]]. The servant ordered to kill them, however, relented and placed the two on the banks of the [[Tiber river]]. The river, which was in flood, rose and gently carried the cradle and the twins downstream, where under the protection of the river deity [[Tiberinus]], they would be adopted by a she-wolf known as ''Lupa'' in [[Latin]], an animal sacred to [[Mars (mythology)|Mars]]. In [[Norse mythology]], [[Fenrir]] or Fenrisulfr is a gigantic wolf, the son of [[Loki]] and the giantess [[Angrboða]]. Fenrir is bound by the gods, but is ultimately destined to grow too large for his bonds and devour [[Odin]] during the course of [[Ragnarök]]. At that time he will have grown so large that his upper jaw touches the sky while his lower touches the earth when he gapes. He will be slain by Odin's son, [[Viðarr]], who will either stab him in the heart or rip his jaws asunder according to different accounts. The [[Bible]] contains 13 references to wolves, usually as metaphors for greed and destructiveness. In the [[New Testament]], [[Jesus]] is quoted to have used wolves as illustrations to the dangers His followers would have faced should they follow him (Matthew 10:16, Acts 10:29, Matthew 7:15)<ref name="BOTF">{{cite book | author= Bright, Michael | title=Beasts of the Field: The Revealing Natural History of Animals in the Bible | year=2006 | pages= p346 | id=ISBN 1861058314 }}</ref> In the medieval Russian folktale [[Ivan Tsarevich and the Grey Wolf]] (also known as ''Ivan Tsarevich and the Firebird'' or ''The Firebird''), the grey wolf is a benevolant character who assists Ivan on his quest, using his knowledge and magical powers. In Japan, grain farmers once worshiped wolves at shrines and left food offerings near their dens, beseeching them to protect their crops from wild boars and deer. Talismans and charms adorned with images of wolves protected against fire, disease, and other calamities and brought fertility to agrarian communities and to couples hoping to have children. The [[Ainu people]] believed that they were born from the union of a wolflike creature and a goddess.<ref name="AINU">{{cite book | author = Walker, Brett L. | title = The Lost Wolves Of Japan | year = 2005 | pages = pp.331 | id = ISBN 0295984929}}</ref> Wolves also figure prominently in the folklore and mythology of some [[Indigenous peoples of the Americas|Native American]] tribes. In the [[Cardinal direction]]s of the [[Plains Indians]], the wolf represented the [[west]], while for the [[Pawnee]], it represented the southeast. According to the Pawnee creation myth, the wolf was the first creature to experience [[death]]. The Wolf Star ([[Sirius]]), enraged at not having been invited to attend a council on how the Earth should be made, sent a wolf to steal the whirlwind bag of The Storm that Comes out of the West, which contained the first humans. Upon being freed from the bag, the humans killed the wolf, thus bringing death into the world. The Pawnee, being both an agricultural and hunting people, associated the wolf with both corn and the bison; the "birth" and "death" of the Wolf Star was to them a reflection of the wolf's coming and going down the path of the [[Milky Way]] known as Wolf Road. Wolves however were not always portrayed positively in Native American cultures. The [[Netsilik Inuit]] and Takanaluk-arnaluk believed that the sea-woman Nuliayuk's home was guarded by wolves. The Naskapi's believed that the [[caribou]] afterlife is guarded by giant wolves which kill careless hunters venturing too near. The [[Navajo people]] feared witches in wolf's clothing called "Mai-cob".<ref name="SAM" /> ===Attacks on humans=== {{Main|Wolf attacks on humans}} [[Image:Wolves of Perigord.jpg|thumb|Two of the [[Wolves of Périgord]], responsible for the deaths of 18 people on February 1766, on display at the chateau of Razac in [[Thiviers]]]] Under normal circumstances, wild wolves are generally timid around humans. Wolves usually try to avoid contact with people, to the point of even abandoning their kills when an approaching human is detected, though there are several reported circumstances in which wolves have been recorded to act aggressively toward humans.<ref name="Attacks">{{cite web | url= http://www.nina.no/archive/nina/Publikasjoner/oppdragsmelding/NINA-OM731.pdf | title= The Fear of Wolves: A Review of Wolf Attacks on Humans | publisher= Norsk Institutt for Naturforskning | accessdate= 2008-06-26|format=PDF}}</ref><ref name="AQ"> {{cite web | url = http://www.aws.vcn.com/wolf_attacks_on_humans.html | title = Wolf attacks on humans | author = Mader, TR | publisher = Abundant Wildlife Society of North America | accessdate = 2007-05-31}}</ref> The frequency with which wolves have been recorded to kill people is much lower than in most other large carnivorous mammals in general, indicating that though potentially dangerous, wolves are among the least threatening for their size and predatory potential.<ref name="Attacks" /> Attacks due to provocation have occured, usually involving shepherds defending their flocks, though none have been recorded to be fatal. Unprovoked attacks by non-rabid wolves are rare, though they have happened. The majority of victims of unprovoked healthy wolves tend to be women and children.<ref name="Attacks" /> Historically, attacks by healthy wolves tended to be clustered in space and time, indicating that human-killing was not a normal behavior for the average wolf, but was rather a specialized behavior that single wolves or packs developed and maintained until they were killed.<ref name="NO" /> Records from the former Soviet Union indicate that the largest number of attacks on children in areas with high wolf densities occurred in summer during July and August, the period when female wolves begin feeding their cubs solid food. Sharp falls in the frequency of attacks were noted in the Autumn months of September and October, coinciding with drops in temperature which caused most children to remain indoors for longer periods.<ref name="Graves"/> [[Habituation]] is a known factor contributing to some wolf attacks which results from living in close proximity to human habitations, which can cause wolves to lose their fear of humans and consequently approach too closely, much like urban coyotes. Habituation can also happen when people intentionally encourage wolves to come up to them, usually by offering them food, or unintentionally, when people do not sufficiently intimidate them.<ref name="Attacks" /> This is corollated by accounts demonstrating that wolves in protected areas are more likely to show [[boldness]] toward humans than ones in areas where they are actively hunted.<ref name="WOLVES">{{cite book | author= L. David Mech & Luigi Boitani | title=Wolves: Behaviour, Ecology and Conservation | year=2001 | pages= p 448 | id= ISBN 0226516962 }}</ref> The majority of fatal wolf attacks have historically involved [[rabies]], which was first recorded in wolves in the 13th century. Though wolves are not reservoirs for the disease, they can catch it from other species. Wolves develop an exceptionally severe aggressive state when infected and can bite numerous people in a single attack. Before a vaccine was developed, bites were almost always fatal. Today, wolf bites can be treated, but the severity of rabid wolf attacks can sometimes result in outright death, or a bite near the head will make the disease act too fast for the treatment to take effect. Unlike healthy wolves which typically limit themselves to attacking women or children, attacks by rabid wolves are made at random, with adult men being killed on occasion. Rabid attacks tend to cluster in winter and spring. With the reduction of rabies in Europe and North America, few rabid wolf attacks have been recorded, though some still occur annually in the middle east.<ref name="Attacks">{{cite web | url= http://www.nina.no/archive/nina/Publikasjoner/oppdragsmelding/NINA-OM731.pdf | title= The Fear of Wolves: A Review of Wolf Attacks on Humans | publisher= Norsk Institutt for Naturforskning | accessdate= 2008-06-26|format=PDF}}</ref> Rabid attacks can be distinguished from predatory attacks by the fact that rabid wolves limit themselves to biting their victims rather than consuming them. Plus, the timespan of predatory attacks can sometimes last for months or years, as opposed to rabid attacks which end usually after a fortnight.<ref name="Attacks" /><ref name="NO">{{cite web | url = http://www.lcie.org/Docs/Regions/Baltic/Linnell%20AZL%20Wolf%20attacks%20in%20Fennoscandia.pdf | title = Is the fear of wolves justified? A Fennoscandian perspective | work = | publisher = Acta Zoologica Lituanica, 2003, Volumen 13, Numerus 1 | accessdate = 2008-05-09|format=PDF}}</ref> Much like some [[big cats]],<ref name="CC">{{cite book | last = Corbett | first = Jim | title = Man-eaters of Kumaon | pages = viii-xiii | publisher = Oxford University Press | year = 1944 }}</ref> old or crippled wolves unable to tackle their normal prey have also been recorded to attack humans.<ref name="Graves"/> People who corner or attack wolves typically receive quick bites to the hands or feet, though the attack is usually not pressed. In both rabid and predatory attacks, victims are usually attacked around the head and neck in a sustained manner,<ref name="Attacks" /> though healthy wolves rarely attack frontally, having been shown to prefer to attack from behind.<ref name="Graves"/> Some specialised [[man-eaters]] have been recorded to kill children by knocking them over from behind and biting the back of their heads and necks.<ref>“Wolves and wild dogs”, from ‘’Man-Eaters’’, Michael Bright, Robson books, 2000</ref> The body of a victim of a healthy wolf is often dragged off and consumed unless disturbed.<ref name="Attacks" /> ===Livestock and pet predation=== [[Image:Chinook2.gif|thumb|''Waiting for a Chinook'', by [[Charles Marion Russell|C.M. Russell]], depicting wolves harassing a steer.]] Wolf depredations on livestock tend to increase in September and October when females teach their cubs how to hunt. Wolves usually attack livestock when they are grazing, though it is not uncommon for some wolves to break into fenced enclosures.<ref name="Graves" /> [[Sheep]] are the most frequently recorded victims in Europe, in India it is [[goat]]s, in Mongolia it is horses, while North American records show wolves having a greater tendency to attack [[cattle]] and [[Turkey (bird)|turkey]]s.<ref name="WOLVES" /> Wolves usually disregard size or age on medium sized prey such as sheep and goats.<ref name="RGWD" /> According to [[Theodore Roosevelt]], the small wolves of the Southern Plains rarely attack full grown cattle or steers, preferring instead young or sick animals, while the large wolves of the northern Rockies can kill fully grown steers unaided.<ref name="TEDDY" /> Injuries may include a crushed skull, severed spine, disembowelment and massive tissue damage. Wolves will also kill sheep by attacking the throat, similar to the manner in which coyotes kill sheep. Wolf kills can be distinguished from coyote kills by the far greater damage to the underlying tissue. Surplus killing often occurs when within the confines of human made livestock shelters.<ref name="RGWD" /> One specimen known as the "Aquila Wolf" in Arizona was known to have killed 65 sheep in one night and 40 at another time.<ref name="Myths">{{cite web | url=http://www.aws.vcn.com/wolf_myth_legend_misconception.html | title = The Wolf: Myth, Legend and Misconception | publisher = Abundant Wildlife Society of North America | accessdate = 2008-07-10}}</ref> Sometimes, the animals survive, but are left with severe mutilations, sometimes warranting [[euthanasia]].<ref name="Myths" /> Livestock with prior experience of a wolf attack may develop behavioural problems, with some animals having been reported to run through [[barbed wire]] fences upon hearing wolves, or refusing to go out into pastures, causing severe weight loss.<ref name="Graves" /> Animals severely injured by wolves often appear dazed and are reluctant to move due to the deep pain.<ref name="RGWD">{{cite web | url=http://animalrangeextension.montana.edu/articles/wildlife/wolf_depredation.htm | title = Ranchers' Guide to Wolf Depredation | publisher = Montana state university | accessdate = 2007-08-24}}</ref> It is often difficult to confirm kills, seeing as wolves often eat all of the animal they have killed if it is below the size of a calf.<ref name="Myths" /> Some non- or less-lethal methods of protecting livestock from wolves have been under development for the past decade. Such methods include [[rubber bullet|rubber]] ammunition and use of guard animals.<ref>{{cite web | url = http://www.geography.wisc.edu/livingwithwolves/reports/2004_Survey_Results_Executive_Summary.pdf | title = Summary Report: 2004/2005 Public Opinion Survey: Wolf Management in Wisconsin | author = Naughton, Lisa, Adrian Treves, Rebecca Grossberg, and David Wilcove | work = Living with Wolves | publisher = UW-Madison Geography | accessdate = 2005-08-30 | format=PDF}}</ref> Recorded wolf howls have also been shown to be effective in at least one incident.<ref name="Ellis">{{cite video|people=[[Shaun Ellis (wolf researcher)|Shaun Ellis]]|title= A Man Among Wolves |medium=[[DVD]]|publisher=[[National Geographic]]|year=2007}}</ref> The extent of livestock losses to wolves vary regionally; from being statistically insignificant, to having critical effects on local economies. In North America, loss of livestock by wolves makes up only a small percentage of total losses. In the [[United States]], wolf predation is low compared to other human or animal sources of livestock loss.<ref>{{cite web | url = http://www.defenders.org/wildlife/new/wolves/loss.html | title = Wolf Predation Plays Small Role in Livestock Losses in 2005 | publisher = Defenders of Wildlife | accessdate = 2006-08-30}}</ref> Since the state of Montana began recording livestock losses due to wolves back in 1987, only 1,200 sheep and cattle have been killed. 1,200 killings in twenty years is not very significant when in the greater Yellowstone region 8,300 cattle and 13,000 sheep die from natural causes. According to the International Wolf Center, a Minnesota-based organization: {{Quotation|To put depredation in perspective, in 1986 the wolf population was at about 1,300–1,400, there were an estimated 232,000 cattle and 16,000 sheep in Minnesota's wolf range. During that year 26 cattle, about 0.01% of the cattle available, and 13 sheep, around 0.08% of the sheep available, were verified as being killed by wolves. Similarly, in 1996 an estimated 68,000 households owned dogs in wolf range and only 10, approximately 0.00015% of the households, experienced wolf depredation.|Wolf Depredation|International Wolf Center, Teaching the World about Wolves<ref>{{ cite web |url = http://www.wolf.org/wolves/learn/intermed/inter_mgmt/depstat.asp | title = Wolf Depredation | publisher = International Wolf Center | date= August 2005}}</ref>}} Furthermore, Jim Dutcher, a film maker who raised a captive wolf pack observed that wolves are very reluctant to try meat that they have not eaten or seen another wolf eat before possibly explaining why [[livestock]] depredation is unlikely except in cases of desperation.<ref>{{cite web | title = Wolves at our door | publisher = Touchstone | first= Jim and Jamie| last= Dutcher | date = 2002| accessdate = 2007-02-27}}</ref> The results however differ in Eurasia. [[Greece]] for example reports that between April 1989 and June 1991, 21000 sheep and goats plus 2729 cattle were killed. In 1998 it was 5894 sheep and goats, 880 cattle and very few horses. <ref name="GW" /> A study on livestock predation taken in [[Tibet]] showed that the wolf was the most prominent predator, accounting for 60% of the total livestock losses, followed by the [[snow leopard]] (38%) and lynx (2%). Goats were the most frequent victims (32%), followed by sheep (30%), yak (15%), and horses (13%). Wolves killed horses significantly more and goats less than would be expected from their relative abundance.<ref>[http://www.springerlink.com/content/42567458h670u205/ "Carnivore-Caused Livestock Mortality in Trans-Himalaya"]</ref> In [[1987]], [[Kazakhstan]] reported over 150,000 domestic livestock losses to wolves, with 200,000 being reported a year later.<ref name="Graves" /> In some areas, [[dog]]s are a major food source for wolves. Reports from Croatia indicate that dogs are killed more frequently than sheep. Wolves in Russia apparently limit feral dog populations. In [[Wisconsin]], more compensation has been paid for dog losses than livestock.<ref name="WOLVES" /> Some wolf pairs have been reported to predate on dogs by having one wolf lure the dog out into heavy brush where the second animal waits in ambush.<ref name="Graves" /> In some instances, wolves have displayed an uncharacteristic fearlessness of humans and buildings when attacking dogs, to an extent where they have to be beaten off or killed.<ref>{{cite web|url= http://www.bbc.co.uk/nature/animals/features/145big.shtml |title=Wolf at my door |accessdate=2007-07-20 |publisher = BBC | accessdate = 2007-08-11}}</ref> Specially bred Livestock guardian dogs have been used to repell wolves from pastures, though their primary function has more to do with intimidating the wolves rather than fighting them.<ref>{{cite web|url=http://www.nal.usda.gov/awic/companimals/guarddogs/guarddogs.htm |title=Livestock Guarding Dogs - Protecting Sheep from Predators |accessdate=2007-07-20 |publisher=[[United States Department of Agriculture]]}}</ref> ===Wolf hunting=== {{Main|Wolf hunting}} [[Image:Wolf-shoot.jpg|A 19th century painting depicting the conclusion of a wolf hunt|thumb]] Wolves are usually hunted for sport, for their skins, to protect [[livestock]], and in some rare cases to protect humans. Wolves are usually hunted in heavy brush and are considered especially challenging to hunt, due to their elusive nature and sharp senses.<ref name="WOLFHUNT">{{cite web | url = http://www.albertabushadventures.com/wolf_hunting.html | title = Alberta Canada Wolf & Coyote Hunts with Alberta Bush Adventures Hunting Guides | work = | publisher = Alberta Bush Adventures| accessdate = 2007-09-27}}</ref> Wolves are notoriously shy and difficult to kill, having been stated to be almost as hard to still hunt as [[cougar]]s, and being far more problematic to dispatch with poison, traps or hounds. Wolves though generally do not defend themselves as effectively as cougars or [[bear]]s.<ref name="TEDDY">{{cite web | url = http://www.fullbooks.com/Hunting-the-Grisly-and-Other-Sketches3.html| title = ''Hunting the Grisly and Other Sketches'' | work = [[Theodore Roosevelt]]| publisher = | accessdate = 2007-09-12}}</ref> Some wolves will evade capture for very long periods of time and display great cunning. On specimen nicknamed "Three Toes of Harding County" in South Dakota eluded its pursuers for 13 years before finally being caught.<ref name="Myths" /> Another wolf nicknamed "[[Rags the digger]]" near Meeker, Colorado would deliberately ruin trap lines by digging up traps without tripping them.<ref name="Outlaw">{{cite web | url = http://www.internationalwolf.net/wolves/news/iwmag/2000/winter/outlaw_wolves.pdf | title = Legends of the "Outlaw Wolves" | work = | publisher = International Wolf Centre| accessdate = 2008-07-12|format=PDF}}</ref> In Sport hunting, wolves are usually taken in late [[Autumn]] and early [[Winter]], when their pelts are of the highest quality and because the heavy snow makes it easier for the wolves to be tracked.<ref name="WOLFHUNT" /> Wolves have occasionally been hunted for food, the meat having been variously described as being tough<ref name="AINU" /> and tasting like [[chicken]].<ref>[http://www.civilization.ca/hist/cae/foo90e.html Civilization.ca - Canadian Arctic Expedition - Food<!-- Bot generated title -->]</ref> The hunting of grey wolves, while originally actively endorsed in many countries, has become a controversial issue in some nations. Opponents see it as [[cruelty to animals|cruel]], unnecessary and based on misconceptions, while proponents argue that it is vital for the [[Conservation movement|conservation]] of game herds and as [[pest control]].<ref name="Prohunt">{{cite web|date=2000|title=Game board says yes to aerial shooting of wolves|publisher= alaskawolves.org|url=http://alaskawolves.org/Blog/DF21F183-DA44-432D-8F73-D9C8DC0BE859_files/Alaska%20Peninsula%20wolf%20control,%20FDNM,%20ADN.pdf|accessdate=2008-04-23|format=PDF}}</ref> ===Reintroduction=== {{main|Wolf reintroduction}} ====North America==== [[Image:Gray wolf endangered species sheet.jpg|thumb|right|Gray wolf endangered species sheet]] In North America, debate about wolf [[reintroduction]] is ongoing and often heated, both where reintroduction is being considered and where it has already occurred. Where wolves have been successfully reintroduced, as in the [[Yellowstone National Park|greater Yellowstone area]] and [[Idaho]], reintroduction opponents continue to cite [[livestock]] predation, [[surplus killing]], and [[economics|economic]] hardships caused by wolves.<ref>{{cite news | url = http://www.tucsonweekly.com/gbase/Currents/Content?oid=oid%3A45520 | first = Tim | last = Vanderpool | title = Politics of the Wolf | publisher = Tucson Weekly|date = [[2002-06-20]] | accessdate = 2006-08-27}}</ref> Opponents in prospective areas echo these same concerns. However, the Yellowstone and Idaho reintroductions demonstrate how compromise can be used to satisfy relevant interests. These reintroductions were the culmination of over two decades of research and [[debate]]. Ultimately, the economic concerns of the local [[ranching]] industry were dealt with when [[Defenders of Wildlife]] decided to establish a fund that would compensate ranchers for livestock lost to wolves, shifting the economic burden from industry to the wolf proponents themselves.<ref>{{cite web | url = http://www.defenders.org/wolfcomp.html | title = The Bailey Wildlife Foundation Wolf Compensation Trust | publisher = Defenders of Wildlife | accessdate = 2006-08-27}}</ref> As of 2005, there are over 450 [[Mackenzie Valley Wolf|Mackenzie Valley wolves]] in the Greater Yellowstone Ecosystem and over 1,000 in Idaho. Both populations have long since met their recovery goals. Lessons learned from this ordeal may yet prove useful where wolf reintroduction continues to create a sharp divide between industry and environmental interests, as it has in [[Arizona]] (where the [[Mexican Wolf]] was released beginning in 1998). Though many hunters, prior to and even after reintroduction, claimed that wolves would wipe out entire populations of [[elk]], [[deer]] and other [[ungulates]], the food chain within the [[Yellowstone]] ecosystem has been re-ordered to deliver a banquet that favors a more varied array of species. Prior to wolf reintroduction, high numbers of elk were linked to declines in aspen and willow communities, which negatively affected beaver and moose. Pre-wolf coyote numbers were much larger, affecting small rodent populations, foxes, and the production of pronghorn antelope. Scavengers had slimmer pickings. Today, with wolves taking elk, reducing their numbers, and leaving more carcasses on the landscape, [[grizzlies]] and [[wolverines]] have easier access to more meat, meaning a better chance for larger litters of cubs and pups. Coyote numbers have been significantly reduced, meaning more mice and pocket gophers for foxes and avian predators like hawks and eagles.<ref>{{cite web| url =http://www.wildlifeconservation.org/ | title= A Brewing Backlash Against Lobos | publisher= Wildlife Conservation | first = Todd | last= Wilkinson | date = March/April 2007| accessdate = 2007-02-27}}</ref> Reports have been published placing the value of revenue from wolf-watching as upward of [[USD|$]]25 million. Native American attitudes toward wolf reintroductions varied. Although the [[Nez Perce]] welcomed the reintroduction of wolves in Idaho, the [[Apache]]s of the southwestern US and [[Kalispell]]s of Washington opposed any reintroduction, as wolves held little spiritual significance in their cultures.<ref name="WOLVES" /> ====United Kingdom==== The British Government signed conventions in the 1980s and 1990s agreeing to consider reintroducing wolves and to promote public awareness about them. Being party to European conventions, the British government is obliged to study the desirability of reintroducing extinct species and to consider reintroducing wolves. Although there are indications that wolves are recolonizing areas in Western Europe, they are unable to return to their former ranges in Britain without active human assistance. The [[Scottish Highlands]] are one of the few large areas in western Europe with a relatively small human population, thus ensuring that wolves would suffer little disturbance from human activity. One popular argument in favour of the reintroduction is that the Highlands' [[red deer]] populations have overgrown. A reintroduction of wolves would aid in keeping their numbers down, thus allowing native flora some respite. Other arguments include the generation of income and local employment in the Highlands through wolf-related ecotourism. This could replace the declining and uneconomical Highland sheep industry. ===Wolves as pets=== Many countries, states and local regions have specific regulations governing the acquisition and management of wolves. In Britain, the keeping of wolves is strictly controlled and a licence is needed to own one. In the United States, the keeping of pure wolves is prohibited by the [[Endangered Species Act Amendments of 1978|U.S. Endangered Species Act]] of 1973. Despite this, there are an estimated 100,000 wolves illegally kept in captivity in the United States, with some surveys concluding that the number could be as high as 2 million.<ref name="WOLVES" /> Captive wolf pups are usually taken from their mother at two weeks of age. Pups will typically develop behavioural problems if raised without another member of their own kind. Due to their wild nature, wolf cubs require more socialisation than dog puppies. Socialising cubs solely with humans runs the risk of the cubs devoting their attention to people rather than their own kind. Wolf pups typically stop responding to socialization at the age of 19 days, as opposed to dogs which can still be socialised at the age of 10 weeks.<ref name="DOGS" /> Due to the fact that wolf milk contains more [[arginine]] than can be found in puppy [[milk substitute]]s, an arginine supplement is needed when feeding pups below the weaning age. Failure to do so can result in the pups developing [[cataracts]].<ref name="WOLVES" /> As adults, wolves have been shown, most of the time, to be largely untrainable and unpredictable, and will sometimes display aggressive behaviour toward small animals and children. Captive wolves are generally shy and avoid [[eye contact]] with humans other than their owner, as well as not listening to any commands made by any other humans. They usually vacate rooms or hide when a new person enters the establishment.<ref>[http://www.wolfsongalaska.org/wolfdogs_characteristics.html Wolf Song of Alaska: Guideline Characteristics of Wolves and Wolfdogs<!-- Bot generated title -->]</ref> Ordinary pet food is inadequate, seeing as an adult wolf needs 1-2.5 kg (2-5 lbs) of quality meat daily along with bones, skin and fur to meet its nutritional requirements. The exercise needs of a wolf exceed the average dog's demand. Because of this, captive wolves typically do not cope well in urban areas. Due to their talent at [[observational learning]], adult captive wolves can quickly work out how to escape confinement<ref name="DOGS" />, and need constant reminding that they are not the leader of their owner/caretaker, which makes raising wolves difficult for people who raise their pets in an even, rather than subordinate, environment. According to the [[American Zoological Association]], the minimum housing recommended for a large canid is an enclosure of 4m x 4m (12 x 12 ft), increased by 50% for each additional canid. To prevent the wolf jumping over the enclosure, fences are specified to be necessarily at least 2m (6 ft) high and needing an overhang at the top. An inside skirt buried below ground is also required to prevent tunnelling. Some pet wolves are [[euthanise]]d or might be released into the wild where they are likely to starve or be killed by resident wolf packs in regions where they are still present. Abandoned or escaped captive wolves can be more destructive and pose a greater danger to humans and livestock than wild wolves, seeing as their habituation to humans causes them to lose their natural shyness.<ref name="Graves" /> The [[Wolf of Gysinge]] is thought to have been one such animal. Captive wolves have also been shown to be unsuitable for working as dogs do. [[German people|German]] wolf biologist [[Erik Zimen]] once attempted to form a [[dog sled]] team composed entirely of pure wolves. The attempt proved to be a complete failure, as the wolves were far more prone to fighting than [[sled dogs]] and ignored most commands.<ref name="DOGS" /> == Media == {{multi-listen start}} {{multi-listen item|filename=Wolf howls.ogg|title=Wolf howl recording|description=|format=[[Ogg]]}} {{multi-listen item|filename=rallying.ogg|title=Rallying|description=European wolves rallying — 157 KB|format=[[Ogg]]}} {{multi-listen end}} == See also == ===Cultural depictions of wolves=== * [[Warg]] * [[Werewolf]] * [[Wolves in fiction]] * [[Wolves in heraldry]] ===Wolf based organizations=== * [[International Wolf Center]] * [[UK Wolf Conservation Trust]] * [[Mission: Wolf]] ===Famous wolves=== *[[Hunting of Jean-Baptiste|Jean-Baptiste]] *[[Lobo the King of Currumpaw]] *[[Rags the digger]] ===Man-eating wolves=== * [[Wolf of Ansbach]] * [[Wolf of Gysinge]] * [[Wolf of Sarlat]] * [[Wolf of Soissons]] * [[Wolves of Ashta]] * [[Wolves of Hazaribagh]] * [[Wolves of Paris]] * [[Wolves of Perigord|Wolves of Périgord]] ===Other extant and extinct canid species also known as wolves=== * [[Falkland Island Wolf]] * [[Maned Wolf]] == Notes and references == {{Reflist|3}} == External links == {{commonscat|Canis lupus}} {{wikispecies|Canis lupus}} {{wikiquote|Wolf}} *[http://www.nwf.org/nationalwildlife/article.cfm?articleId=829&issueId=69 Return of gray wolf to Yellowstone park] *[http://www.wolf.org The International Wolf Center] {{Canidae nav}} {{North American Game}} {{Heraldic creatures}} {{DEFAULTSORT:Wolf, Gray}} [[Category:Arctic land animals]] [[Category:Fauna of Finland]] [[Category:Fauna of Ireland]] [[Category:Fauna of Russia]] [[Category:Fauna of the Middle East]] [[Category:Mammals of Asia]] [[Category:Mammals of Canada]] [[Category:Mammals of Europe]] [[Category:Mammals of India]] [[Category:Mammals of North America]] [[Category:Mammals of the United States]] [[Category:Wolves|Gray Wolf]] {{Link FA|ast}} {{Link FA|de}} {{Link FA|ka}} {{Link FA|no}} {{Link FA|pl}} {{Link FA|uk}} [[ang:Wulf]] [[ar:ذئب رمادي]] [[an:Lupo]] [[frp:Lop]] [[ast:Llobu]] [[zh-min-nan:Lông]] [[ba:Бүре]] [[bar:Wuif]] [[bs:Vuk]] [[br:Bleiz gris]] [[bg:Вълк]] [[ca:Llop]] [[cv:Кашкăр]] [[cs:Vlk obecný]] [[co:Lupu]] [[cy:Blaidd]] [[da:Ulv]] [[de:Wolf]] [[nv:Ma’iitsoh]] [[et:Hunt]] [[el:Λύκος]] [[eml:Lauv]] [[es:Canis lupus]] [[eo:Lupo]] [[eu:Otso]] [[fa:گرگ]] [[fo:Úlvur]] [[fr:Loup]] [[gd:Faol]] [[gl:Lobo]] [[got:𐍅𐌿𐌻𐍆𐍃]] [[hak:Lòng]] [[ko:늑대]] [[hr:Sivi vuk]] [[io:Volfo]] [[id:Serigala abu-abu]] [[ia:Lupo gris]] [[is:Úlfur]] [[it:Canis lupus]] [[he:זאב מצוי]] [[ka:მგელი]] [[kk:Қасқыр]] [[ky:Карышкыр]] [[ht:Lou]] [[ku:Gur]] [[la:Lupus]] [[lv:Vilks]] [[lt:Pilkasis vilkas]] [[lij:Canis lupus]] [[lmo:Lop]] [[hu:Farkas]] [[nah:Cuetlāchtli]] [[myv:Верьгиз]] [[nl:Wolf (dier)]] [[cr:ᒪᐦᐃᐦᑲᓐ]] [[ja:オオカミ]] [[no:Gråulv]] [[nn:Ulv]] [[nrm:Loup]] [[oc:Canis lupus]] [[pl:Wilk]] [[pt:Lobo]] [[ro:Lup cenuşiu]] [[qu:Lupu]] [[ru:Волк]] [[sah:Бөрө]] [[scn:Lupu]] [[simple:Wolf]] [[sk:Vlk dravý]] [[cu:Влькъ]] [[sl:Sivi volk]] [[sr:Вук]] [[sh:Vuk]] [[fi:Susi]] [[szl:Wilk]] [[sv:Varg]] [[tl:Lobo (hayop)]] [[ta:ஓநாய்]] [[th:หมาป่า]] [[vi:Chó sói xám]] [[tg:Гург]] [[tr:Kurt]] [[uk:Вовк звичайний]] [[fiu-vro:Susi]] [[wa:Leu]] [[bat-smg:Vėlks]] [[zh:狼]]