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Overview

Brief Summary

Description

"Gray wolves usually live in packs, led by an ""alpha pair."" The pack includes some of the alpha pair's offspring and may include some unrelated wolves. A pack's territory can be as large as 13,000 square km. Howling probably helps advertise who ""owns"" a particular piece of territory. When pups are born, the mother stays near them for the first three weeks, and her mate and others in the pack bring food to her. The pups are weaned when they about nine weeks old. As adults, they may travel as far as 72 km a day with their pack and run as fast as 70 km per hour."

Adaptation: This transparent view of the wolf braincase shows how the brain is situated within. The large feature projecting from the front of the brain is the olfactory lobe, an important center related to the sense of smell.

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  • Original description: Linnaeus, C., 1758.  Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth Edition. Laurentii Salvii, Stockholm, 1:39, 824 pp.
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Comprehensive Description

Longueur : 110-150 cm dont 30 à 50 cm de queue. Taille au garrot : 60-90 cm. Poids : 20-40 kg pour le mâle et 18-30 kg pour la femelle.

La silhouette générale du Loup ressemble à celle d’un chien de berger mais avec un avant-train plus puissant. Sa tête est plus large et les oreilles sont proportionnellement plus petites, plus écartées et de forme générale plutôt arrondie. Ses yeux sont obliques avec un iris jaune. Il possède 5 doigts aux pattes avant et 4 aux pattes arrières et son empreinte ressemble à s’y méprendre à celle d’un grand chien. En France, le Loup a un pelage beige-gris avec un liseré noir sur les pattes antérieures. Le bout de la queue est noir. Le dos peut avoir des nuances plus foncées par rapport au ventre. Il présente un masque facial clair. Comme pour la plupart des canidés, le loup compte 42 dents : I3/3, C1/1, P4/4, M2/3. Les prémolaires P4 supérieures et les molaires M1 inférieures sont modifiées en carnassières permettant au carnivore de broyer les os et de découper la chair des proies.

Social, le Loup vit en meute, composée de 2 à 8 individus en France, dont la composition se renouvelle régulièrement. La hiérarchie de la meute est très stricte. Ainsi, elle est régie par un couple dominante (alpha), suivi d’un mâle bêta, puis des subordonnées. Le couple alpha dirige les activités vitales (chasse, déplacements, défense du territoire et reproduction). Seul ce couple se reproduit. La maturité sexuelle est atteinte à l’âge de 2 ans. L’accouplement a lieu en février-mars. La femelle met bas en avril-mai, après une gestation de 62-63 jours, de 3 à 5 louveteaux aveugles et sourds et pesant environ 400-500 g. Ils sont sevrés vers 2 mois et peuvent vivre jusqu’à 14-16 ans.

Le Loup est un carnivore opportuniste. Il adapte donc son régime aux proies disponibles : cerfs, chamois, mouflons, chevreuils, sangliers. Il peut aussi se nourrir de lièvres, rongeurs ou oiseaux et peut s’attaquer aux ovins domestiques.

Territorial, son domaine vital s’étend sur 150 à 300 km2 en France. Ubiquiste, il peut se trouver dans des milieux très variés allant des plaines, steppes et savanes jusqu’en montagnes. Son domaine vital possède au moins une tanière, lieu indispensable pour la reproduction car c’est là que la femelle va mettre bas et élever les jeunes. La tanière peut-être une cavité creusée par le Loup lui-même, une tanière de Renard ou de Blaireau agrandie, une grotte, une cavité sous une souche ou un rocher, dans un taillis très dense… et doit être située à proximité d’un point d’eau.

Références :
BEAUFORT (DE) F. 1987. Le Loup en France : élément d’écologie historique. Société française pour l’Etude et la Protection des Mammifères. Encyclopédie des Carnivores de France 1 : 32p.
ONCFS. 1998-2012. Quoi de neuf ? Bulletin du réseau Loup 1-27. Disponible sur : http://www.oncfs.gouv.fr
ONCFS. 2011. Le Loup. Disponible sur : http://www.oncfs.gouv.fr
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Description of Canis familiaris

The domestic dog. This is regarded by most people as a subspecies of Canis lupus, see Canis lupus for a description.
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Description of Canis lupus

There are about 40 suspecies of Canis lupus (dogs and wolves) , including two domesticated subspecies (Canis lupus dingo in Australia and Canis lupus familiaris - mans best friend). Many subspecies of wolf occur throughout the Northern hemisphere. Canis lupus is assessed as Least Concern by the IUCN, as its relatively widespread range and stable population trend mean that the species, at global level, does not meet, or nearly meet, any of the criteria for the threatened categories. However, some local populations are classified as Endangered and some subspecies are endangered or extinct. The domestic dogs were derived from the gray wolf domestication about 15,000 years ago. The dog may have been the first animal to be domesticated, and has been the most widely kept working, hunting, and companion animal in human history. They quickly became widespread across world cultures. Dogs perform many roles for people, such as hunting, herding, pulling loads, protection, assisting police and military, companionship, and, more recently, aiding handicapped individuals. In some cultures, dogs are also an important source of meat. In 2001, there were estimated to be 400 million dogs in the world. Over the 15,000-year span in which the dog has been domesticated, it has diverged into a handful of landraces, groups of similar animals whose morphology and behavior have been shaped by environmental factors and functional roles.Selective breeding has produced hundreds of varied breeds, showing more behavioral and morphological variation than any other land mammal. Height measured to the withers ranges from a few inches in the Chihuahua to a few feet in the Irish Wolfhound; color varies from white through grays (usually called blue) to black, and browns from light (tan) to dark (red or chocolate) in a wide variation of patterns.
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Distribution

occurs (regularly, as a native taxon) in multiple nations

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National Distribution

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) Formerly throughout North America south through much of Mexico; also Europe and Asia. Replaced by the red wolf in the southeastern United States. Today found south of Canada only in northern Mexico (no recent confirmed reports; extirpated or maybe a few in eastern Sonora, Chihuahua, and/or Zacatecas?), a few areas in the Rocky Mountains (northwestern Montana, reintroduction sites in Wyoming and Idaho), northwestern Great Lakes region (northeastern third of Minnesota, northern Wisconsin, Michigan Upper Peninsula), and Cascade Mountains of northern Washington. Formerly much more numerous in the Rocky Mountain states than in the southwestern U.S. (Johnson 1991). Extirpated in much of southern Canada (see Theberge [1992] and Can. Field-Nat. 106:138 for range/status map); remains in 85% of former total Canadian range (Theberge 1991).

In 1995, wolf reintroductions were initiated in the Yellowstone ecosystem and in central Idaho (nonessential experimental populations) (USFWS 1994; Federal Register, 16 August 1994; Bangs and Fritts 1993; End. Sp. Bull. 20(4):4-5). See Bangs et al. (1998) for information on the status of gray wolf restoration in Montana, Idaho, and Wyoming. In 1998, USFWS (Federal Register, 12 January 1998) announced its intention to reintroduce the Mexican gray wolf (subspecies baileyi) into Arizona and New Mexico (Apache and Gila national forests, also possibly White Sands Missile Range).

Wolf observations in the Dakotas have increased in recent years, likely related to range expansion and population increases in adjacent areas, especially Minnesota; most occurrences have been of young individuals, suggesting dispersal (Licht and Fritts 1994).

Grewal et al. (2004) used genetic data to determine that the wolf population in Algonquin Provincial Park in Ontario, Canada, is a southern part of a larger metapopulation of Canis lycaon (or Canis lupus lycaon).

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Range Description

Originally, the Grey Wolf was the world's most widely distributed mammal, living throughout the northern hemisphere north of 15°N latitude in North America and 12°N in India. It has become extinct in much of Western Europe (Boitani 1995), in Mexico and much of the USA (Mech 1970). Their present distribution is more restricted: wolves occur primarily in wilderness and remote areas, especially in Canada, Alaska and northern USA, Europe, and Asia from about 75°N to 12°N.
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Historic Range:
Holarctic

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Geographic Range

Gray wolves once ranged throughout the Northern hemisphere, from the Arctic south to southern Mexico, northern Africa, and southern Asia. However, due to habitat destruction, environmental change, persecution by humans, and other barriers to population growth, gray wolf populations are now found only in a few areas of the contiguous United States, Alaska, Canada, Mexico (a small population), and Eurasia.

Biogeographic Regions: nearctic (Native ); palearctic (Native )

Other Geographic Terms: holarctic

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The gray wolf is found worldwide, ranging from across Europe to northern
Asia; however, it has been extirpated from much of its former range.
Formerly in North America, the gray wolf ranged from the southern fringe of
Greenland south through mid-Mexico and from the Atlantic to the Pacific
[34].  It occupied almost all regions of the United States except for
deserts and high mountaintops [22,34].  Today the gray wolf occupies about 1
percent of its former range in the contiguous states [10].  It occupies
northeastern Minnesota, northern Wisconsin, the Upper Peninsula of
Michigan, northwestern Montana, central Idaho, and Washington's Cascade
Mountains.  In addition the gray wolf is abundant throughout Alaska and
Canada.  The ranges for the 24 subspecies follow [22,34]:

Ssp. irremotus - Idaho, western Montana, Wyoming, Alberta, and the
                 western fringes of Washington and Oregon
Ssp. columbianus - British Columbia and southwestern Alberta; can
                   move into the northwestern states
Ssp. occidentalis -  northern Alberta and Saskatchewan,
                     northeastern British Columbia, and central
                     Manitoba, into the Yukon and the Northwest
                     Territories
Ssp. lycaon - southeastern Manitoba, Ontario, Quebec, and the
              eastern United States, from the Atlantic to
              central Minnesota, south to northeastern Florida
Ssp. nubilus - thought to be extinct, although it may possibly
               occur in Minnesota [19]; from southern Manitoba
               and Saskatchewan, south through the Great Plains
               into northern Texas
Ssp. alces - the Kenai Peninsula, Alaska
Ssp. pambasileus - Yukon Territory and all but northern Alaska
Ssp. tundrarum - northern Alaska
Ssp. hudsonicus - along the Hudson Bay in the Northwest
                  Territories and Manitoba
Ssp. arctos - Melville Island, Northwest Territories
Ssp. orion - Greenland
Ssp. labradorius - northern Quebec and Newfoundland
Ssp. beothucus - the island of Newfoundland
Ssp. ligoni - Alexander Archipelago, Alaska
Ssp. fuscus - the Cascade Mountains of Washington, Oregon, and
              California
Ssp. crassodon - Vancouver Island, British Columbia
Ssp. youngi - the southern Rocky Mountains of Utah, Arizona,
              New Mexico, Colorado, and Wyoming
Ssp. mogollonensis - central Arizona and westcentral New Mexico
Ssp. monstrabilis - Texas, Mexico, and southeast New Mexico
Ssp. baileyi - central Mexico into southern Arizona and
               New Mexico
Ssp. bernardi - Banks and Victoria Islands, Northwest Territories
Ssp. mackenzii - northern Northwest Territories and Yukon Territory
Ssp. manningi - Baffin Island, Northwest Territories
Ssp. griseoalbus - Alberta, Saskatchewan, Manitoba, Northwest
                   Territories, and Newfoundland
  • 10. Fritts, Steven H.; Mech, L. David. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf population in northwest Minnesota. Wildlife Monographs No. 80. Washington, DC: The Wildlife Society. 79 p. [13863]
  • 19. Mech, L. David; Frenzel, L. D., Jr. 1971. The possible occurrence of the great plains wolf in northeastern Minnesota. In: Mech, L. David; Frenzel, L. D., Jr., eds. Ecological studies of the timber wolf in northeastern Minnesota. Res. Pap. NC-52. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 60-62. [13891]
  • 22. Mech, L. David. 1974. Canis lupus. Mammalian Species. 37: 1-6. [13866]
  • 34. Young, Stanley P.; Goldman, Edward A. 1944. The wolves of North America: parts I and II. New York: Dover Publishers. 636 p. [13861]

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Regional Distribution in the Western United States

More info on this topic.

This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

    2  Cascade Mountains
    8  Northern Rocky Mountains
    9  Middle Rocky Mountains
   16  Upper Missouri Basin and Broken Lands

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Occurrence in North America

AK ID MI MN MT WA WV WI WY
AB BC MB NB NF NT NS ON PE PQ
SK YT

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Geographic Range

The original range of Canis lupus consisted of the majority of the Northern hemisphere -- from the Arctic continuing south to a latitude of 20° S, which runs through southern Central Mexico, northern Africa, and southern Asia. However, due to habitat destruction, environmental change, persecution by humans, and other barriers to population growth, gray wolf populations are now found only in a few areas of the contiguous United States, Alaska, Canada, Mexico (a small population), and Eurasia.

Biogeographic Regions: nearctic (Native ); palearctic (Native )

Other Geographic Terms: holarctic

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Distribution in Egypt

Localized? (South Sinai).

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Physical Description

Morphology

Physical Description

Gray wolves are the largest wild dog species in the world. They vary in size depending on where they are found, with southern populations being generally smaller than northern populations. Total body length, from tip of the nose to tip of the tail, is from 870 to 1,300 mm. Height (measured from base of paws to shoulder) generally ranges from 60 to 90 cm. Males are, on average, larger than females.

Fur color of gray wolves also varies depending on where they are living. Their color ranges from pure white in Arctic populations, to mixtures of white with gray, brown, cinammon, and black to nearly uniform black in some color phases. Their upperparts are generally darker and they have lighter fur on their undersides, the tail is tipped with black.

Gray wolves have a dense underfur layer, providing them with excellent insulation against cold conditions.

Range mass: 23.0 to 80 kg.

Range length: 870 to 1300 mm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

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Physical Description

The largest of approximately 41 wild species of canids, gray wolves vary in size based primarily on geographic locality, with southern populations generally smaller than northern populations. Total body length, from tip of the nose to tip of the tail, is from 1000 to 1300 mm in males, and 870 to 1170 mm in females. Tail length ranges between 350 to 520 mm. Males can weigh from 30 to 80 kg, with an average of 55 kg, females can weigh from 23 to 55 kg, with an average of 45 kg. Height (measured from base of paws to shoulder) generally ranges from 60 to 90 cm. Distance between the canines is around 4 cm.

Fur color of gray wolves also varies geographically, ranging from pure white in Arctic populations, to mixtures of white with gray, brown, cinammon, and black to nearly uniform black in some color phases.

North American populations have three distinct color phases. The normal phase is characterized by varying mixtures of white with shades of black, gray, cinnamon, and brown on the upper parts of the animal. The back is usually more profoundly black, and the muzzle, ears, and limbs have cinammon coloration as well. Under parts are whitish and the tail is conspicuously black over the tail gland, and paler below to the tip, which is nearly pure black. The black phase of North American populations is characterized by the upper parts varying from brown to black, with specks of white; the underparts are paler in tone, and there is often a pure white medial pectoral spot. The third color phase occurs during the first pelage of young wolves. The upper parts are drab-gray, overlaid with brownish-black. The underparts are paler as well, and the ears vary from black to buffy, depending on the subspecies (Young 1944).

Gray wolves have a dense underfur layer, providing them with excellent insulation against cold conditions.

Gray wolves can be distinguished from red wolves (Canis rufus) by their larger size, broader snout, and shorter ears. They are distinguished from coyotes (Canis latrans) by being 50 to 100% larger and having a broader snout and larger feet.

Range mass: 23.0 to 80 kg.

Range length: 870 to 1300 mm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

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Size

Length: 205 cm

Weight: 40000 grams

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Size in North America

Length:
Average: 1,100 mm males; 1,050 kg, females
Range: 1,000-1,300 mm males; 870-1,170 mm females

Weight:
Average: 55 kg males; 45 kg females
Range: 30-80 kg males; 23-55 kg females
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Diagnostic Description

Differs from the coyote in larger nose pad (greater than 1 inch vs. less than 1 inch), more rounded ears, larger anteroposterior diameter of upper canine at gum level (more than 11 mm vs. less than 11 mm), larger heel pad on forefoot (more than 1.25 inches vs. less than 1.25 inches), longer skull (more than 215 mm and 2.5 mm vs. less than 215 mm and 205 mm, for males and females, respectively), and relatively shorter canines (in coyote, tips of upper canines extend below level of anterior mental foramina when lower jaw is in place); also, gray wolf holds the tail high when running, coyote holds it low (see Hall 1981 and Hoffmeister 1986 for further details). In some parts of central and eastern North America, coyote approaches wolf in certain characteristics, due to interbreeding. Differs from the red wolf in larger size, longer skull (condylobasal length more than 213 mm and 203 mm vs. less than these measurements for males and females, respectively), and in certain features og the molars (see Hall 1981); however, the red wolf actually may be a coyote-gray wolf hybrid (see GTAXCOM for CANIS RUFUS). Differs from the domestic dog in generally larger size, broader nose pad, more massive skull with heavier teeth, rostrum relatively longer, supraoccipital shield larger and projecting farther posteriorly, front foot track longer and narrower (Hoffmeister 1986).

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Ecology

Habitat

Palouse Grasslands Habitat

This taxon is found in the Palouse grasslands, among other North American ecoregions. The Palouse ecoregion extends over eastern Washington, northwestern Idaho and northeastern Oregon. Grasslands and savannas once covered extensive areas of the inter-mountain west, from southwest Canada into western Montana in the USA. Today, areas like the great Palouse prairie of eastern  are virtually eliminated as natural areas due to conversion to rangeland. The Palouse, formerly a vast expanse of native wheatgrasses (Agropyron spp), Idaho Fescue (Festuca idahoensis), and other grasses, has been mostly plowed and converted to wheat fields or is covered by Drooping Brome (Bromus tectorum) and other alien plant species.

the Palouse historically resembled the mixed-grass vegetation of the Central grasslands, except for the absence of short grasses. Such species as Bluebunch Wheatgrass (Elymus spicatus), Idaho Fescue (Festuca idahoensis) and Giant Wildrye (Elymus condensatus) and the associated species Lassen County Bluegrass (Poa limosa), Crested Hairgrass (Koeleria pyramidata), Bottlebrush Squirrel-tail (Sitanion hystrix), Needle-and-thread (Stipa comata) and Western Wheatgrass (Agropyron smithii) historically dominated the Palouse prairie grassland.

Representative mammals found in the Palouse grasslands include the Yellow-bellied Marmot (Marmota flaviventris), found burrowing in grasslands or beneath rocky scree; American Black Bear (Ursus americanus); American Pika (Ochotona princeps); Coast Mole (Scapanus orarius), who consumes chiefly earthworms and insects; Golden-mantled Ground Squirrel (Spermophilus lateralis); Gray Wolf (Canis lupus); Great Basin Pocket Mouse (Perognathus parvus); Northern River Otter (Lontra canadensis); the Near Threatened Washington Ground Squirrel (Spermophilus washingtoni), a taxon who prefers habitat with dense grass cover and deep soils; and the Northern Flying Squirrel (Glaucomys sabrinus), a mammal that can be either arboreal or fossorial.

There are not a large number of amphibians in this ecoregion. The species present are the Great Basin Spadefoot Toad (Spea intermontana), a fossorial toad that sometimes filches the burrows of small mammals; Long-toed Salamander (Ambystoma macrodactylum); Northern Leopard Frog (Glaucomys sabrinus), typically found near permanent water bodies or marsh; Columbia Spotted Frog (Rana luteiventris), usually found near permanent lotic water; Pacific Treefrog (Pseudacris regilla), who deposits eggs on submerged plant stems or the bottom of water bodies; Tiger Salamander (Ambystoma tigrinum), fossorial species found in burrows or under rocks; Woodhouse's Toad (Anaxyrus woodhousii), found in arid grasslands with deep friable soils; Western Toad (Anaxyrus boreas), who uses woody debris or submerged vegetation to protect its egg-masses.

There are a limited number of reptiles found in the Palouse grasslands, namely only: the Northern Alligator Lizard (Elgaria coerulea), often found in screes, rock outcrops as well as riparian vicinity; the Painted Turtle (Chrysemys picta), who prefers lentic freshwater habitat with a thick mud layer; Yellow-bellied Racer (Chrysemys picta); Ringneck Snake (Diadophis punctatus), often found under loose stones in this ecoregion; Pygmy Short-horned Lizard (Phrynosoma douglasii), a fossorial taxon often found in bunchgrass habitats; Side-blotched Lizard (Uta stansburiana), frequently found in sandy washes with scattered rocks; Southern Alligator Lizard (Elgaria multicarinata), an essentially terrestrial species that prefers riparian areas and other moist habitats; Pacific Pond Turtle (Emys marmorata), a species that usually overwinters in upland habitat; Western Rattlesnake (Crotalus viridis), who, when inactive, may hide under rocks or in animal burrows; Night Snake (Hypsiglena torquata); Western Skink (Plestiodon skiltonianus), who prefers grasslands with rocky areas; Western Terrestrial Garter Snake (Thamnophis elegans), found in rocky grasslands, especially near water; Rubber Boa (Charina bottae).

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Arizona Mountains Forests Habitat

This taxon is found in the Arizona Mountain Forests, which extend from the Kaibab Plateau in northern Arizona to south of the Mogollon Plateau into portions of southwestern Mexico and eastern Arizona, USA. The species richness in this ecoregion is moderate, with vertebrate taxa numbering 375 species. The topography consists chiefly of steep foothills and mountains, but includes some deeply dissected high plateaus. Soil types have not been well defined; however, most soils are entisols, with alfisols and inceptisols in upland areas. Stony terrain and rock outcrops occupy large areas on the mountains and foothills.

The Transition Zone in this region (1980 to 2440 m in elevation) comprises a strong Mexican fasciation, including Chihuahua Pine (Pinus leiophylla) and Apache Pine (P. engelmannii) and unique varieties of Ponderosa Pine (P. ponderosa var. arizonica). Such forests are open and park-like and contain many bird species from Mexico seldom seen in the U.S.. The Canadian Zone (above 2000 m) includes mostly Rocky Mountain species of mixed-conifer communities such as Douglas-fir (Pseudotsuga menziesii), Engelmann Spruce (Picea engelmanni), Subalpine Fir (Abies lasiocarpa), and Corkbark Fir (A. lasiocarpa var. arizonica). Dwarf Juniper (Juniperus communis) is an understory shrubby closely associated with spruce/fir forests. Exposed sites include Chihuahua White Pine (Pinus strobiformis), while disturbed north-facing sites consists primarily of Lodgepole Pine (Pinus contorta) or Quaking Aspen (Populus tremuloides).

There are a variety of mammalian species found in this ecoregion, including the endemic Arizona Gray Squirrel (Sciurus arizonensis), an herbivore who feeds on a wide spectrum of berries, bark and other vegetable material. Non-endemic mammals occurring in the ecoregion include: the Banner-tailed Kangaroo Rat (Dipodomys spectabilis NT); Desert Pocket Gopher (Geomys arenarius NT). In addition, there is great potential for restoring Mexican Wolf (Canis lupus) and Grizzly Bear (Ursus arctos horribilis) populations in the area because of its remoteness and juxtaposition to other ecoregions where these species were formerly prevalent.

There are few amphibians found in the Arizona mountain forests. Anuran species occurring here are: Red-spotted Toad (Anaxyrus punctatus); Southwestern Toad (Anaxyrus microscaphus); New Mexico Spadefoot Toad (Spea multiplicata); Woodhouse's Toad (Anaxyrus woodhousii); Northern Leopard Frog (Lithobates pipiens); Chiricahua Leopard Frog (Lithobates chiricahuensis VU); Madrean Treefrog (Hyla eximia), a montane anuran found at the northern limit of its range in this ecoregion; Boreal Chorus Frog (Anaxyrus woodhousii); Western Chorus Frog (Pseudacris triseriata); and Canyon Treefrog (Hyla arenicolor). The Jemez Mountains Salamander (Plethodon neomexicanus NT) is an ecoregion endemic, found only in the Jemez Mountains of Los Alamos and Sandoval counties, New Mexico. Another salamander occurring in the ecoregion is the Tiger Salamander (Ambystoma tigrinum).

A number of reptilian taxa occur in the Arizona mountains forests, including: Gila Monster (Heloderma suspectum NT), often associated with cacti or desert scrub type vegetation; Narrow-headed Garter Snake (Thamnophis rufipunctatus), a near-endemic found chiefly in the Mogollon Rim area; Sonoran Mud Turtle (Kinosternon sonoriense NT).

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Comments: No particular habitat preference. In Minnesota and Wisconsin, usually occurs in areas with few roads, which increase human access and incompatible land uses (Thiel 1985, Mech et al. 1988, Mech 1989) but apparently can occupy semi-wild lands if ungulate prey are abundant and if not killed by humans (see Mladenoff et al. 1997). Minimum of 10,000-13,000 sq km (with low road density) might be necessary to support a viable population (USFWS 1990); a single pack does not constitute a "minimum viable population" (USFWS 1990). Young are born in an underground burrow that has been abandoned by another mammal or dug by wolf. In Northwest Territories, dens were most commonly located witin 50 km of northern tree line, which resulted in maximal availability of caribou during the denning and pup rearing period; within the tundra zone, dens were not preferentially located near caribou calving grounds (Heard and Williams 1992). In Minnesota, dens usually were not near territory boundaries; den use was traditional in most denning alpha females studied for more than 1 year; possibly the availability of a stable food supply source helped determine den location (Ciucci and Mech 1992).

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Habitat and Ecology

Habitat and Ecology
Ranges in all northern habitats where there is suitable food (Mech 1970), densities being highest where prey biomass is highest (Fuller 1989). Food is extremely variable, but the majority comprises large ungulates (moose, caribou, deer, elk, wild boar, etc.). Wolves will also eat smaller prey items, livestock, carrion, and garbage.

Systems
  • Terrestrial
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Gray wolves are one of the most wide ranging land animals. They occupy a wide variety of habitats, from arctic tundra to forest, prairie, and arid landscapes.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: tundra ; taiga ; chaparral ; forest ; scrub forest ; mountains

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Cover Requirements

More info for the term: tree

Gray wolves excavate natal dens in well-drained soils in meadows near water
[16].  They may use the same den for several years.  In Minnesota Fuller
[11] found gray wolves denning in hollow logs (24 to 35 inches [60-90 cm]
diameter).  Gray wolves also den under tree roots, rock outcrops, or even in
beaver lodges [11].  After 1 to 2 months these natal dens are abandoned
for an open area called a rendez-vous site.  Here the pups are guarded
by a few adult pack members, while the rest of the pack hunts [1].
Herman and Willard [16] summarized that gray wolves need a large, remote area
relatively free from human disturbance.  Territory sizes range from 20
to 215 square miles (54-555 sq km) in Minnesota [10].  Average territory
sizes in Minnesota have been reported to vary from 55 to 120 square
miles (143-310 sq km) [29] and 25 to 29 square miles (64-75 sq km) [2].
In the West average territory sizes vary from 75 to 150 square miles
(194-388 sq km) and are smaller in winter when ungulates are
concentrated on their wintering grounds [16].
  • 1. Ballard, Warren B.; Whitman, Jackson S.; Gardner, Craig L. 1987. Ecology of an exploited wolf population in south-central Alaska. Wildlife Monographs No. 98. Washington, DC: The Wildlife Society. 54 p. [13865]
  • 2. Berg, William E.; Kuehn, David W. 1982. Ecology of wolves in northeast Minnesota. In: Harrington, Fred H.; Paquet, Paul C., eds. Wolves of the world: Perspectives of behavior, ecology, and conservation. Park Ridge, NJ: Noyes Publications: 4-11. [13868]
  • 10. Fritts, Steven H.; Mech, L. David. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf population in northwest Minnesota. Wildlife Monographs No. 80. Washington, DC: The Wildlife Society. 79 p. [13863]
  • 11. Fuller, Todd K. 1989. Population dynamics of wolves in northcentral Minnesota. Wildlife Monographs No. 105. Washington, DC: The Wildlife Society. 41 p. [13864]
  • 16. Herman, Margaret, Willard, E. Earl. 1978. Rocky Mountain wolf and its habitat. Missoula, MT: U.S. Department of Agriculture, Forest Service, National Forest System Cooperative Forestry, Forestry Research, Region 1. 17 p. [16522]
  • 29. Scott, Barbara M. V.; Shackleton, David M. 1982. A preliminary study of the social organization of the Vancouver Island wolf. In: Harrington, Fred H.; Paquet, Paul C., eds. Wolves of the world: Perspectives of behavior, ecology, and conservation. Park Ridge, NJ: Noyes Publications: 12-24. [13869]

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Preferred Habitat

Gray wolves' habitat preferences appear to be more prey dependent than cover
dependent.  Herman and Willard [16] summarized that gray wolves choose home
territories with a variety of topographic features.  Forests, open
meadows, rocky ridges, and lakes or rivers all comprise a pack's
territory.  In the West gray wolves have been known to follow the seasonal
elevational movements of ungulate herds [16].  In Minnesota, where
territories encompass only subtle elevational changes, Fritts and Mech
[10] observed no changes in territory use by gray wolves between summer and
winter.  In south-central Alaska Ballard and others [1] found that
gray wolves do not follow migrating moose or caribou outside of their pack
territories.  Gray wolves do, however, follow moose and caribous' elevational
movements within pack territories.
  • 1. Ballard, Warren B.; Whitman, Jackson S.; Gardner, Craig L. 1987. Ecology of an exploited wolf population in south-central Alaska. Wildlife Monographs No. 98. Washington, DC: The Wildlife Society. 54 p. [13865]
  • 10. Fritts, Steven H.; Mech, L. David. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf population in northwest Minnesota. Wildlife Monographs No. 80. Washington, DC: The Wildlife Society. 79 p. [13863]
  • 16. Herman, Margaret, Willard, E. Earl. 1978. Rocky Mountain wolf and its habitat. Missoula, MT: U.S. Department of Agriculture, Forest Service, National Forest System Cooperative Forestry, Forestry Research, Region 1. 17 p. [16522]

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Associated Plant Communities

More info for the term: tundra

Gray wolves inhabit a variety of plant communities.  Their territories
usually contain a mix of forested and open areas.  Gray wolves can also be
found on the tundra.  In the West, gray wolves have been known to follow
ungulate herds from their lowland wintering grounds to their high summer
pastures [16].  In the East, gray wolves
inhabit a mix of coniferous and
deciduous forests, which include balsam fir (Abies balsamea), black
spruce (Picea mariana), white spruce (P. glauca), white-cedar (Thuja
occidentalis), jack pine (Pinus banksiana), white pine (P. strobus), red
pine (P. resinosa), tamarack (Larix laricina), sugar maple (Acer
saccharum), yellow birch (Betula alleghaniensis), and eastern hemlock
(Tsuga canadensis).  In the West, gray wolves inhabit Douglas-fir (Pseudotsuga
menziesii)-spruce (Picea spp.) forests, as well as ponderosa pine (Pinus
ponderosa) and western larch (Larix occidentalis) forests [16,23,28].
  • 16. Herman, Margaret, Willard, E. Earl. 1978. Rocky Mountain wolf and its habitat. Missoula, MT: U.S. Department of Agriculture, Forest Service, National Forest System Cooperative Forestry, Forestry Research, Region 1. 17 p. [16522]
  • 23. Mech, L. David; Karns, Patrick D. 1977. Role of the wolf in a deer decline in the Superior National Forest. Res. Pap. NC-148. St. Paul, MN: U.S. Department of Agriculture, Forest Service,North Central Forest Experiment Station. 23 p. [13886]
  • 28. Robinson, William L.; Werner, J. Kirwin. 1975. Vertebrate animal populations of the McCormick Forest. Res. Pap. NC-118. St. Paul, MN: U.S. Department of Agriculture, Forest Service,North Central Forest Experiment Station. 25 p. [13484]

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Habitat: Cover Types

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This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

     1  Jack pine
     5  Balsam fir
    12  Black spruce
    13  Black spruce - tamarack
    15  Red pine
    16  Aspen
    17  Pin cherry
    18  Paper birch
    20  White pine - northern red oak - red maple
    21  Eastern white pine
    24  Hemlock - yellow birch
    25  Sugar maple - beech - yellow birch
    30  Red spruce - yellow birch
    33  Red spruce - balsam fir
    35  Paper birch - red spruce - balsam fir
    37  Northern white cedar
    38  Tamarack
   107  White spruce
   108  Red maple
   201  White spruce
   202  White spruce - paper birch
   203  Balsam poplar
   204  Black spruce
   205  Mountain hemlock
   206  Engelmann spruce - subalpine fir
   208  Whitebark pine
   210  Interior Douglas-fir
   211  White fir
   212  Western larch
   213  Grand fir
   215  Western white pine
   217  Aspen
   218  Lodgepole pine
   220  Rocky Mountain juniper
   221  Red alder
   222  Black cottonwood - willow
   223  Sitka spruce
   224  Western hemlock
   225  Western hemlock - Sitka spruce
   230  Douglas-fir - western hemlock
   235  Cottonwood - willow
   237  Interior ponderosa pine
   253  Black spruce - white spruce
   254  Black spruce - paper birch

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Habitat: Plant Associations

More info on this topic.

This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the term: shrub

   K003  Silver fir - Douglas-fir forest
   K004  Fir - hemlock forest
   K008  Lodgepole pine - subalpine forest
   K010  Ponderosa shrub forest
   K011  Western ponderosa forest
   K012  Douglas-fir forest
   K013  Cedar - hemlock - pine forest
   K014  Grand fir - Douglas-fir forest
   K015  Western spruce - fir forest
   K093  Great Lakes spruce - fir forest
   K094  Conifer bog
   K095  Great Lakes pine forest
   K096  Northeastern spruce - fir forest
   K106  Northern hardwoods
   K107  Northern hardwoods - fir forest

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Habitat: Ecosystem

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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

FRES10 White-red-jack pine
FRES11 Spruce-fir
FRES18 Maple-beech-birch
FRES19 Aspen-birch
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES22 Western white pine
FRES23 Fir-spruce
FRES24 Hemlock-Sitka spruce
FRES25 Larch
FRES26 Lodgepole pine
FRES36 Mountain grasslands
FRES37 Mountain meadows

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Gray wolves are one of the most wide ranging land animals. They occupy a wide variety of habitats, from arctic tundra to forest, prairie, and arid landscapes.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: tundra ; taiga ; chaparral ; forest ; scrub forest ; mountains

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Migration

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

Wolves in far northwestern North America may move as required to remain with migratory caribou.

Home ranges very large but very variable as well, generally ranging from less 100 to 10,000s of square kilometers. In Minnesota, Fritts and Mech (1981) found territory sizes ranging from 195 to 555 square kilometers; in south to central Alaska, Ballard et al. (1987) reported territory sizes from 943 to 2541 square kilometers; in the southern Yukon, Hayes (1992) found territory sizes of 583 to 794 km square kilometers; in the Kenai Peninsula, Alaska, Peterson et al. (1984) found average territory sizes of 638 square kilometers; and in coastal forests of Vancouver Island, Atkinson and Janz (1994) found territory sizes from 100 to 400 square kilometers. Packs that depend on barren ground caribou migrate with the caribou as far as 360 km (Kuyt 1972; Mech 1970, 1974). In the Glacier National Park area, territory size averages around 780 sq km (Bangs and Fritts 1993).

In the Northwest Territories and western Nunavut, Walton et al. (2001) fitted 23 wolves in 19 different packs with collar-mounted satellite transmitters. Annual home-range sizes (95% minimum convex polygon) averaged 63,058 sq km for males and 44,936 sq km for females. Straight-line distances from the most distant location on the winter range to the den site averaged 508 km in 1997-1998 and 265 km in 1998-1999 (wolves followed caribou). All but 2 of 15 wolves returned to within 25 km of a previous den, and 2 wolves returned to the same den site.

Dispersing young may move several hundred kilometers. In Minnesota, most dispersers left when they were 11-12 months old; dispersal occurred mainly in February-April and October-November; 35% of known-age wolves remained in their natal territory for more than 2 years (Gese and Mech 1991).

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Trophic Strategy

Comments: Predominant prey: ungulates. When these are low or seasonally unavailable, eats alternative prey, such as beaver, snowshoe hare, rodents, and carrion. Commonly hunts in packs, but lone wolves and pairs are able to kill prey as large as adult moose (Thurber and Peterson 1993). In the vicinity of Glacier National Park, feeds primarily on white-tailed deer; sometimes kills mountain lions and sometimes usurps ungulate prey killed by lions (Bangs and Fritts 1993). White-tailed deer and moose carrion were the primary prey in southern Ontario (Forbes and Theberge 1996).

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Food Habits

Gray wolves are carnivores. They hunt prey on their own, in packs, steal the prey of other predators, or scavenge carrion. Prey is located by chance or by scent. What wolves eat depends on where they live and what kinds of prey are available. Wolves hunt in packs for large prey such as Alces alces, Cervus elaphus, Bison bison, musk oxen, and Rangifer tarandus. Wolves help to keep prey populations healthy by hunting the weak, old, and sick animals. A wolf can consume up to 9 kg of meat at one meal. Wolves usually eat almost the entire carcass, including some hair and bones. Smaller animals, such as Castor canadensis, Sylvilagus floridanus, and other small mammals are usually hunted by lone wolves. Wolves may also eat livestock and garbage when it is available.

Animal Foods: mammals; carrion

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Food Habits

Gray wolves prey mainly on large ungulates, such as moose (Alces alces), deer
(Odocoileus spp.), elk (Cervus elaphus), and caribou (Rangifer
tarandus).  They tend to prey on the young, old, and sick members of
ungulate populations.  Beaver (Castor canadensis) are a major supplement
to gray wolves' diets [23].  Voigt and others [33] reported that gray wolves'
diets vary, depending on relative prey abundance.  Other prey species
include mountain goats (Oreamnos americanus), bison (Bison bison),
pronghorn (Antilocapra americana), various rodents, upland game birds
and waterfowl, snowshoe hare (Lepus americanus), and black bear (Ursus
americana) [6,10,21,23,25,33]. On Isle Royale seeds of wild sarsaparilla
(Aralia nudicaulis) were found in gray wolf scat [7].  Occasionally gray wolves
prey on domestic livestock.
  • 6. Cohn, Jeffrey P. 1990. Endangered wolf population increases. Bioscience. 40(9): 628-632. [13887]
  • 7. Edwards, Joan. 1985. Effects of herbivory by moose on flower and fruit production of Aralia nudicaulis. Journal of Ecology. 73: 861-868. [13626]
  • 10. Fritts, Steven H.; Mech, L. David. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf population in northwest Minnesota. Wildlife Monographs No. 80. Washington, DC: The Wildlife Society. 79 p. [13863]
  • 21. Mech, L. David. 1973. Wolf numbers in the Superior National Forest of Minnesota. Res. Pap. NC-97. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 10 p. [13885]
  • 23. Mech, L. David; Karns, Patrick D. 1977. Role of the wolf in a deer decline in the Superior National Forest. Res. Pap. NC-148. St. Paul, MN: U.S. Department of Agriculture, Forest Service,North Central Forest Experiment Station. 23 p. [13886]
  • 25. Meehan, William R. 1974. The forest ecosystem of southeast Alaska: 4. Wildlife habitats. Gen. Tech. Rep. PNW-16. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 32 p. [13479]
  • 33. Voigt, Dennis R.; Kolenosky, George B.; Pimlott, Douglas H. 1976. Changes in summer foods of wolves in central Ontario. Journal of Wildlife Management. 40(4): 663-668. [13862]

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Food Habits

Gray wolves are carnivores. They hunt prey on their own, in packs, steal the prey of other predators, or scavenge carrion. Prey is located by chance or scent. Animals included in the diet of gray wolves varies geographically and depends on prey availability. Wolves primarily hunt in packs for large prey such as moose, elk, bison, musk oxen, and reindeer. Once these large ungulates are taken down, the wolves attack their rump, flank, and shoulder areas. Wolves control prey populations by hunting the weak, old, and immature. A wolf can consume up to 9 kg of meat at one meal. Wolves usually utilize the entire carcass, including some hair and bones. Smaller prey such as beavers, rabbits, and other small mammals are usually hunted by lone wolves, and they are a substantial part of their diet. Wolves may also eat livestock and garbage when it is available.

Animal Foods: mammals; carrion

Primary Diet: carnivore (Eats terrestrial vertebrates)

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Associations

Ecosystem Roles

As top predators, gray wolves are important in regulating populations of their prey animals.

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Predation

Few animals prey on gray wolves. Wolves and coyotes are highly territorial animals so wolves from other packs and Canis latrans will attack wolves that are alone or young. They will kill pups if they find them.

Known Predators:

  • coyotes (Canis_latrans)
  • gray wolves (Canis_lupus)

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Predators

Humans are the only significant predator of the gray wolf and have eradicated
it from almost all of its former range worldwide [27,34].  Pimlott and
others [26] noted black bear preying on gray wolf cubs and adults.
  • 26. Pimlott, D. H.; Shannon, J. A.; Kolenosky, G. B. 1969. The ecology of the timber wolf in Algonquin Provincial Park. Ottawa, ON: Candian Forestry Service, Department of Lands and Forests. 91 p. [13904]
  • 27. Ream, Robert R.; Mattson, Ursula I. 1982. Wolf status in the northern Rockies. In: Harrington, Fred H.; Paquet, Paul C., eds. Wolves of the world: Perspectives of behavior, ecology, and conservation. Park Ridge, NJ: Noyes Publications: 362-381. [13871]
  • 34. Young, Stanley P.; Goldman, Edward A. 1944. The wolves of North America: parts I and II. New York: Dover Publishers. 636 p. [13861]

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Ecosystem Roles

As top predators, gray wolves are important in regulating populations of their prey animals.

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Predation

Few animals prey on gray wolves. Wolves and coyotes are highly territorial animals so wolves from other packs and coyotes will attack wolves that are alone or young. They will kill pups if they find them.

Known Predators:

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Known predators

Canis lupus is prey of:
Canis lupus
Canis latrans

This list may not be complete but is based on published studies.
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Known prey organisms

  • L. W. Swan, The ecology of the high Himalayas, Sci. Am. 205:68-78, from pp. 76-77 (October 1961).
  • I. K. Sharma, A study of ecosystems of the Indian desert, Trans. Indian Soc. Desert Technol. and Univ. Center Desert Stud. 5(2):51-55, from p. 52 and A study of agro-ecosystems in the Indian desert, ibid. 5:77-82, from p. 79 1980).
  • Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2006. The Animal Diversity Web (online). Accessed February 16, 2011 at http://animaldiversity.org. http://www.animaldiversity.org
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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 81 to >300

Comments: Because wolves are wide-ranging it is difficult to estimate the number of distinct occurrences. The 30-km separation between EOs given in EOSPECS is a compromise figure. It could have been smaller in the northern USA where pack home ranges in Minnesota were 130 km2, or larger in Alaska where home ranges up to 13,000 km2 are reported (Mech and Frenzel 1971, Burkholder 1959).

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Global Abundance

10,000 to >1,000,000 individuals

Comments: Canada: about 50,000-58,500 in 1990; subject to hunting and trapping in all but a few percent of the range (Cohn 1990, Theberge 1991).

Alaska: about 6000 (Theberge 1991).

In 1998, there were at least 65 wolves in northwestern Montana, at least 122 in central Idaho, and 116 in the Greater Yellowstone Area (Bangs et al. 1998). In 2004, the northern Rocky Mountains population included more than 800 wolves (USFWS 2006).

Isle Royale population was about 14-16 in 1995. Michigan Upper Peninsula population was 112 in 1996/1997 (End. Sp. Bull. 22(3):21). Wisconsin: 150 in 1997-1997 (End. Sp. Bull. 22(3):21). Minnesota: nearly 2000 in mid-1990s (Mladenoff et al. 1997).

Southwestern U.S. and Mexico: none in the U.S. and very few if any in Mexico.

See Jhala and Giles (1991) for information on status in northwestern India (several hundred remain). Population estimates for the 1980s: former Soviet Union, 88,000; Inner Mongolia, less than 10,000; Yugoslavia, 2000-5000; Poland, about 1000; Spain, about 1000; 2000 or less in each of Iran, Afganistan, India, and Romania (see Theberge 1991). Population in the USSR in the mid-1940s was about 200,000-300,000; reduced to lowest numbers in history by the 1960s; increased to 1940s level by the 1980s; about 40,000 in Russia in 1995 (Yuli Gabur 1996, Natural Areas News 1(2):8-9).

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General Ecology

Territorial throughout the year in most areas (but see Migration/Mobility comments). Packs consist of one or more family groups (generally 2-8 members, up to 21) with dominance hierarchy. In the Glacier National Park area, packs generally include 8-12 individuals (Bangs and Fritts 1993). Not uncommonly solitary; lone wolves may move through territories of established packs (e.g., see Thurber and Peterson 1993).

Population density is low; at Isle Royale National Park, Michigan, peak density was 9/100 sq km. Population density on Isle Royale followed trends in food supply (moose), with lag of 2-3 years (Peterson and Page 1988).

Generally wolves are not instrumental in causing prey declines; effect varies with other circumstances. In Quebec, winter weather appeared to affect deer population trend more than did wolf predation (Potvin et al. 1992). In south-central Alaska, wolf predation may have limited caribou recruitment (Bergerud and Ballard 1988), though winter starvation also was proposed as a significant poplation control. May take livestock as secondary prey when deer fawns (the primary summer prey) are less vulnerable due to better prenatal nutrition resulting from mild winter (USFWS 1990). In Minnesota, snow-induced changes in deer distribution and mobility resulted in changes in wolf movement patterns, sociality, and feeding behavior (when snow was shallow, wolves traveled farther and more often, spent less time with pack members, and used conifer cover less and killed fewer deer there) (Fuller 1991).

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Timing of Major Life History Events

Mating - occurs from January to April
Gestation Period - 63 days
Litter Size - average five to six pups; weaned at 5 weeks
Breeding Age - 2 years, but often do not breed until 3 years due to
               social structure of the pack; usually only dominant
               male and female breed
Life Span - up to 16 years, but 10 years is considered quite old
Pack Size - averages 2 to 15 individuals, although 36 individuals have
            been reported; packs structured in a dominance hierarchy
[10,21,22,35]
  • 10. Fritts, Steven H.; Mech, L. David. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf population in northwest Minnesota. Wildlife Monographs No. 80. Washington, DC: The Wildlife Society. 79 p. [13863]
  • 21. Mech, L. David. 1973. Wolf numbers in the Superior National Forest of Minnesota. Res. Pap. NC-97. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 10 p. [13885]
  • 22. Mech, L. David. 1974. Canis lupus. Mammalian Species. 37: 1-6. [13866]
  • 35. Zeveloff, Samuel I.; Collett, Farrell R. 1988. Mammals of the Intermountain West. Salt Lake, UT: University of Utah Press. 365 p. [13860]

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Habitat-related Fire Effects

The effect of fire on gray wolf habitat is best defined by how fire affects
gray wolves' prey.  Beaver, elk, moose, and deer are fire-dependent species,
requiring the plant communities that persist following frequent fires
[14,17].  Edwards [8] reported that after fire moose populated the area
around Wells Gray Park, British Columbia, where they were previously
unknown.  This was followed by a marked increase in gray wolves.  Other
studies in Alaska, Michigan, Minnesota, and Canada show an increase in
moose populations following fire [14,15,32].

Now absent from the old-growth forests of Minnesota, caribou once were
an important prey for gray wolves here.  These forests do not provide enough
food to sustain other ungulates for gray wolves to prey on.  Due to fire
exclusion, these old-growth forested areas have increased, checking
ungulate populations and consequently limiting gray wolf populations [15].
  • 8. Edwards, Joan. 1985. Effects of herbivory by moose on flower and fruit production of Aralia nudicaulis. Journal of Ecology. 73: 861-868. [13626]
  • 14. Hansen, H. L.; Krefting, L. W.; Kurmis, V. 1973. The forest of Isle Royale in relation to fire history and wildlife. Tech. Bull. 294; Forestry Series 13. Minneapolis, MN: University of Minnesota, Agricultural Experiment Station. 44 p. [8120]
  • 15. Heinselman, Miron L. 1973. Fire in the virgin forests of the Boundary Waters Canoe Area, Minnesota. Quaternary Research. 3: 329-382. [282]
  • 17. Kramp, Betty A.; Patton, David R.; Brady, Ward W. 1983. The effects of fire on wildlife habitat and species. RUN WILD: Wildlife/ habitat relationships. Albuerque, NM: U.S. Department of Agriculture, Forest Service, Southwestern Region, Wildlife Unit Technical Report. 29 p. [152]
  • 32. Viereck, Leslie A.; Schandelmeier, Linda A. 1980. Effects of fire in Alaska and adjacent Canada--a literature review. BLM-Alaska Tech. Rep. 6. Anchorage, AK: U.S. Department of the Interior, Bureau of Land Mangement, Alaska State Office. 124 p. [7075]

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Life History and Behavior

Behavior

Communication and Perception

Rank is communicated among wolves by body language and facial expressions, such as crouching, chin touching, and rolling over to show their stomach.

Vocalizations, such as howling allows pack members to communicate with each other about where they are, when they should assemble for group hunts, and to communicate with other packs about where the boundaries of their territories are. Scent marking is ordinarily only done by the alpha male, and is used for communication with other packs.

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Communication and Perception

Rank is communicated among wolves by body language and facial expressions, such as crouching, chin touching, and rolling over to show their stomach.

Vocalizations, such as howling allows pack members to communicate with each other about where they are, when they should assemble for group hunts, and to communicate with other packs about where the boundaries of their territories are. Scent marking is ordinarily only done by the alpha male, and is used for communication with other packs.

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Cyclicity

Comments: Mainly nocturnal. In fall-winter in Minnesota, spends most of time sleeping, resting, or traveling, little time feeding (Mech, 1992, J. Mamm. 73:570-571). In south-central Alaska, den site activity was mainly nocturnal; there was a high probability that groups of wolves would be present at the den at midday (Ballard et al., 1991, Can. Field-Nat. 105:497-504).

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Life Expectancy

Lifespan/Longevity

Gray wolves may live thirteen years in the wild, though average lifespan is 5 to 6 years.  As adults they usually die from old age or from injuries received while hunting or fighting with other wolves.  In captivity they may live to be fifteen years of age.

Range lifespan

Status: wild:
13.0 (high) years.

Average lifespan

Status: wild:
5.0 years.

Range lifespan

Status: captivity:
15.0 (high) years.

Average lifespan

Status: wild:
10.0 years.

Average lifespan

Status: captivity:
20.0 years.

Average lifespan

Sex: female

Status: wild:
13.7 years.

Average lifespan

Sex: female

Status: captivity:
16.0 years.

Average lifespan

Status: wild:
16.0 years.

Average lifespan

Status: captivity:
20.0 years.

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Lifespan/Longevity

Gray wolves may live thirteen years in the wild, though average lifespan is 5 to 6 years.  As adults they usually die from old age or from injuries received while hunting or fighting with other wolves.  In captivity they may live to be fifteen years of age.

Range lifespan

Status: wild:
13.0 (high) years.

Average lifespan

Status: wild:
5.0 years.

Range lifespan

Status: captivity:
15.0 (high) years.

Average lifespan

Status: wild:
10.0 years.

Average lifespan

Status: captivity:
20.0 years.

Average lifespan

Sex: female

Status: wild:
13.7 years.

Average lifespan

Sex: female

Status: captivity:
16.0 years.

Average lifespan

Status: wild:
16.0 years.

Average lifespan

Status: captivity:
20.0 years.

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Lifespan, longevity, and ageing

Maximum longevity: 20.6 years (captivity) Observations: In the wild, 10 year-old females have been known to give birth and record longevity is 13.7 years (Ronald Nowak 1999). One wild born male was around 20.6 years old when he died in captivity (Richard Weigl 2005).
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Reproduction

Breeds late fall/early winter in south, February-March in north. Gestation lasts about 2 months. Young are born in March and early April in the south (Hoffmeister 1986), late April in northwestern Montana, late May-early June in Northwest Territories (Heard and Williams 1992). Litter size is 4-10 (average 6-7); 1 litter/year. Only the dominant male/female mate and rear offspring. Pups emerge from the den in about 3 weeks. Pups are weaned in 50 days (also reported as 5 weeks). Young and parents vacate the den when young are about 3 months old (Hoffmeister 1986). Some offspring remain with the pack, others disperse as they mature. Breeding first occurs in the second or third year (Hoffmeister 1986). Lone wolves generally do not successfully rear young, but they may if food is abundant (Boyd and Jimenez, 1994, J. Mamm. 75:14-17).

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The dominant pair in a grey wolf pack are the only members that breed. This pair is monogamous although, with the death of an alpha individual, a new alpha male or female will emerge and take over as the mate.

Mating System: monogamous ; cooperative breeder

Breeding occurs between the months of January and March, with northern populations breeding later in the season than southern populations. Female gray wolves choose their mates and often form a life-long pair bond. Gray wolf pairs spend a great deal of time together. After mating occurs, the female digs a den in which to raise her young. The den is often dug with an entrance that slopes down and then up again to a higher area to avoid flooding. Pups are born in the den and will remain there for several weeks after birth. Other dens are under cliffs, under fallen trees, and in caves.

The gestation period lasts between 60 and 63 days, litter size ranges from five to fourteen, with the average size being seven pups.

Female pups reach maturity at two years of age, while males will not reach full maturity until three years of age.

Breeding interval: Gray wolves breed once each year.

Breeding season: Gray wolves breed between January and March, depending on where they are living.

Range number of offspring: 5.0 to 14.0.

Average number of offspring: 7.0.

Range gestation period: 63.0 (high) days.

Average weaning age: 45.0 days.

Range age at sexual or reproductive maturity (female): 2.0 to 3.0 years.

Range age at sexual or reproductive maturity (male): 2.0 to 3.0 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); fertilization ; viviparous

Average birth mass: 450 g.

Average number of offspring: 6.

Gray wolf pups are born blind and deaf. They weigh approximately 0.5 kg and depend on the mother for warmth. At ten to fifteen days of age, the pups' blue eyes open, but they only have control over their front legs, thus crawling is their only mode of mobility. Five to ten days later, the young are able to stand, walk, and vocalize.  Pups are cared for by all members of the pack. Until they are 45 days old the pups are fed regurgitated food by all pack members. They are fed meat provided by pack members after that age. During the 20th to 77th day, the pups leave the den for the first time and learn to play fight. Interactions at this time, as well as the dominance status of the mother, ultimately determines their position in the pack hierarchy. Wolf pups develop rapidly, they must be large and accomplished enough to hunt with the pack with the onset of winter. At approximately ten months old, the young begin to hunt with the pack.

Parental Investment: no parental involvement; altricial ; pre-fertilization (Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Male, Female, Protecting: Male, Female); pre-independence (Provisioning: Male, Female, Protecting: Male, Female); post-independence association with parents; extended period of juvenile learning; maternal position in the dominance hierarchy affects status of young

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The dominant pair in a grey wolf pack are the only members that breed. This pair is monogamous although, with the death of an alpha individual, a new alpha male or female will emerge and take over as the mate.

Mating System: monogamous ; cooperative breeder

Breeding occurs between the months of January and April, with northern populations breeding later in the season than southern populations. Female gray wolves choose their mates and often form a life-long pair bond. Gray wolf pairs spend a great deal of time together. Female gray wolves come into estrus once each year and lasts 5 to 14 days, mating occurs during this time. After mating occurs, the female digs a den in which to raise her young. The den is often dug with an entrance that slopes down and then up again to a higher area to avoid flooding. Pups are born in the den and will remain there for several weeks after birth. Other dens are under cliffs, under fallen trees, and in caves.

The gestation period lasts between 60 and 63 days, litter size ranges from one to fourteen, with the average size being six or seven pups. Pups remain in the den until they are 8 to 10 weeks old. Females stay with their pups almost exclusively for the first 3 weeks. Pups are cared for by all members of the pack. Until they are 45 days old the pups are fed regurgitated food by all pack members. They are fed meat provided by pack members after that age.

Female pups reach maturity at two years of age, while males will not reach full maturity until three years of age. Most young gray wolves disperse from their natal pack when they are between 1 and 3 years old.

Breeding interval: Gray wolves breed once each year.

Breeding season: Gray wolves breed between January and March, depending on where they are living.

Range number of offspring: 5.0 to 14.0.

Average number of offspring: 7.0.

Range gestation period: 63.0 (high) days.

Average weaning age: 45.0 days.

Range age at sexual or reproductive maturity (female): 2.0 to 3.0 years.

Range age at sexual or reproductive maturity (male): 2.0 to 3.0 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); fertilization ; viviparous

Average birth mass: 450 g.

Average number of offspring: 6.

Gray wolf pups are born blind and deaf. They weigh approximately 0.5 kg and depend on the mother for warmth. At ten to fifteen days of age, the pups' blue eyes open, but they only have control over their front legs, thus crawling is their only mode of mobility. Five to ten days later, the young are able to stand, walk, and vocalize.  Pups are cared for by all members of the pack. Until they are 45 days old the pups are fed regurgitated food by all pack members. They are fed meat provided by pack members after that age. During the 20th to 77th day, the pups leave the den for the first time and learn to play fight. Interactions at this time, as well as the dominance status of the mother, ultimately determines their position in the pack hierarchy. Wolf pups develop rapidly, they must be large and accomplished enough to hunt with the pack with the onset of winter. At approximately ten months old, the young begin to hunt with the pack.

Parental Investment: no parental involvement; altricial ; pre-fertilization (Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Male, Female, Protecting: Male, Female); pre-independence (Provisioning: Male, Female, Protecting: Male, Female); post-independence association with parents; extended period of juvenile learning; maternal position in the dominance hierarchy affects status of young

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Evolution and Systematics

Functional Adaptations

Functional adaptation

Senses detect epileptic seizures: dog
 

Dogs can determine when their owners are about to have an epileptic seizure by sensing subtle cues.

   
  "The British Epilepsy Association believes that epilepsy-prescient dogs such as these are sensing two very subtle clues to impending epilepsy attacks. One is a high-pitched sound emitted by epileptics up to half an hour before an attack takes place, which is thought to be linked to the abnormal electrical impulses in the brain that occur prior to a seizure. The other is an exceedingly faint odor emitted by epileptics at about the same time as the sound, whose origin is as yet unexplained. Both clues cannot be detected by humans, but can be discerned by the dogs' more acute senses of hearing and smell." (Shuker 2001:226)
  Learn more about this functional adaptation.
  • Shuker, KPN. 2001. The Hidden Powers of Animals: Uncovering the Secrets of Nature. London: Marshall Editions Ltd. 240 p.
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Functional adaptation

Nose sniffs out cancer: dog
 

The noses of some domestic dogs can detect some forms of cancer in humans via an acute sense of smell.

   
  "In the course of some bibliographical research during the early 1990s, Dr. Armand Cognetta, a dermatologist based in Florida, was surprised to discover in the medical literature a number of confirmed cases in which patients had been found to possess hitherto-unsuspected skin cancers that were detected after their pet dogs (usually for several months before the diagnosis) had been compulsively sniffing the area of skin containing the malignancy. Indeed, in each case it had been the behavior of the dog that had finally prompted the owner to seek medical advice in order to find out why their pet was acting so strangely." (Shuker 2001:227)
  Learn more about this functional adaptation.
  • Shuker, KPN. 2001. The Hidden Powers of Animals: Uncovering the Secrets of Nature. London: Marshall Editions Ltd. 240 p.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage: Canis lupus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 271
Specimens with Barcodes: 283
Species With Barcodes: 1
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Barcode data: Canis familiaris

The following is a representative barcode sequence, the centroid of all available sequences for this species.


No available public DNA sequences.

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Statistics of barcoding coverage: Canis familiaris

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 1
Specimens with Barcodes: 31
Species With Barcodes: 1
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Barcode data: Canis lupus

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 284 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

AACCGATGATTGTTCTCCACTAATCACAAGGATATTGGTACTTTATACTTACTATTTGGAGCATGAGCCGGTATAGTAGGCACTGCCTTG---AGCCTCCTCATCCGAGCCGAACTAGGTCAGCCCGGTACTTTACTAGGTGAC---GATCAAATTTATAATGTCATCGTAACCGCCCATGCTTTCGTAATAATCTTCTTCATAGTCATGCCCATCATAATTGGGGGCTTTGGAAACTGACTAGTGCCGTTAATA---ATTGGTGCTCCGGACATGGCATTCCCCCGAATAAATAACATGAGCTTCTGACTCCTTCCTCCATCCTTTCTTCTACTATTAGCATCTTCTATGGTAGAAGCAGGTGCAGGAACGGGATGAACCGTATACCCCCCACTGGCTGGCAATCTGGCCCATGCAGGAGCATCCGTTGACCTT---ACAATTTTCTCCTTACACTTAGCCGGAGTCTCTTCTATTTTAGGGGCAATTAATTTCATCACTACTATTATCAACATAAAACCCCCTGCAATATCCCAGTATCAAACTCCCCTGTTTGTATGATCAGTACTAATTACAGCAGTTCTACTCTTACTATCCCTGCCTGTACTGGCTGCT---GGAATTACAATACTTTTAACAGACCGGAATCTTAATACAACATTTTTTGATCCCGCTGGAGGAGGAGACCCTATCCTATATCAACACCTATTCTGATTCTTCGGGCATCCTGAAGTTTACATTCTTATCCTGCCCGGATTCGGAATAATTTCTCACATTGTCACTTACTACTCAGGGAAAAAA---GAGCCTTTCGGTTATATAGGAATAGTATGAGCAATAATATCTATTGGGTTTTTAGGCTTTATCGTATGAGCTCACCATATGTTTACCGTAGGAATAGATGTAGACACACGAGCATACTTTACGTCCGCCACTATAATTATCGCTATTCCAACGGGAGTAAAAGTATTTAGTTGACTG---GCAACACTTCATGGAGGC---AATATTAAATGATCTCCAGCTATGCTATGAGCTTTAGGGTTTATTTTCTTATTTACAGTAGGCGGGTTAACAGGTATTGTCCTAGCTAATTCGTCCTTAGACATCGTTCTTCATGATACATATTATGTTGTAGCTCATTTTCACTATGTG---CTTTCAATAGGAGCAGTTTTTGCCATTATGGGAGGATTTGCCCACTGATTCCCTTTATTCTCAGGTTATACTCTTAACGATACTTGAGCAAAGATTCACTTTACAATTATGTTTGTGGGAGTAAATATAACTTTCTTCCCTCAACATTTCCTAGGTTTATCTGGAATACCTCGT---CGATACTCTGACTACCCAGATGCATATACT---ACCTGAAATACCGTCTCCTCTATAGGATCGTTTATCTCGCTTACAGCGGTGATGCTTATAATTTTTATGATTTGGGAAGCCTTTGCATCCAAACGAGAAGTT---GCTATAGTAGAACTTACTACAACTAAC
-- end --

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Conservation

Conservation Status

National NatureServe Conservation Status

Canada

Rounded National Status Rank: N5 - Secure

United States

Rounded National Status Rank: N4 - Apparently Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G4 - Apparently Secure

Reasons: Holarctic distribution; survives in wilderness that is not subject to human population pressures; extirpated from most of contiguous U.S. due to human-caused direct mortality; reintroduced populations in Yellowstone and central Idaho have been increasing rapidly; many (tens of thousands) remain in Canada/Alaska, about 2000 south of Canada, 100,000+ in Palearctic. NatureServe rank calculator version 3.1 yielded a rank of G5? and this species is nationally ranked N5 in Canada. However, until a more comprehensive review can be done by NatureServe, our rank will remain G4G5.

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IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2010

Assessor/s
Mech, L.D. & Boitani, L. (IUCN SSC Wolf Specialist Group)

Reviewer/s
Sillero-Zubiri, C. & Hoffmann, M.

Contributor/s

Justification

Originally, the Grey Wolf was the world's most widely distributed mammal. It has become extinct in much of Western Europe, in Mexico and much of the USA, and their present distribution is more restricted; wolves occur primarily but not exclusively in wilderness and remote areas. Their original worldwide range has been reduced by about one-third by deliberate persecution due to depredation on livestock and fear of attacks on humans. Since about 1970, legal protection, land-use changes and rural human population shifts to cities have arrested wolf population declines and fostered natural recolonization in parts of its range and reintroduction in three areas of USA. Continued threats include competition with humans for livestock and game species, exaggerated concern by the public regarding the threat and danger of wolves, and fragmentation of habitat, with resulting areas becoming too small for populations with long-term viability.

Although the Grey Wolf still faces some threats, its relatively widespread range and stable population trend mean that the species, at global level, does not meet, or nearly meet, any of the criteria for the threatened categories. Therefore, it is assessed as Least Concern (LC). However, at regional level, several wolf populations are seriously threatened. In North America, some of the reintroduced populations are still threatened; and in Europe, http://www.iucnredlist.org/apps/redlist/details/3746/1, http://www.iucnredlist.org/apps/redlist/details/3746/3, the species is classified as LC globally but several regional populations, such as the Western-Central Alps population, are classified as Endangered (http://www.lcie.org/).


History
  • 2004
    Least Concern
  • 2004
    Least Concern
    (IUCN 2004)
  • 1994
    Vulnerable
    (Groombridge 1994)
  • 1990
    Vulnerable
    (IUCN 1990)
  • 1988
    Vulnerable
    (IUCN Conservation Monitoring Centre 1988)
  • 1986
    Vulnerable
    (IUCN Conservation Monitoring Centre 1986)
  • 1982
    Vulnerable
    (Thornback and Jenkins 1982)
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Current Listing Status Summary

Status: Endangered
Date Listed: 03/11/1967
Lead Region:   Mountain-Prairie Region (Region 6)   
Where Listed: U.S.A.: All of AL, AR, CA, CO, CT, DE, FL, GA, KS, KY, LA, MA, MD, ME, MO, MS, NC, NE, NH, NJ, NV, NY, OK, PA, RI, SC, TN, VA, VT and WV; those portions of AZ, NM, and TX not included in an experimental population; and portions of IA, IN, IL, ND, OH, OR, SD, UT, and WA. Mexico.

Status: Experimental Population, Non-Essential
Date Listed: 01/12/1998
Lead Region:   Southwest Region (Region 2)   
Where Listed: Mexican gray wolf, EXPN population

Status: Delisted due to Recovery
Date Listed: 01/27/2012
Lead Region:   Great Lakes-Big Rivers Region (Region 3)   
Where Listed: Western Great Lakes DPS

Status: Delisted due to Recovery
Date Listed: 05/05/2011
Lead Region:   Mountain-Prairie Region (Region 6)   
Where Listed: Northern Rocky Mountain DPS (delisted, except WY)

Status: Delisted due to Recovery
Date Listed: 05/05/2011
Lead Region:   Mountain-Prairie Region (Region 6)   
Where Listed: WY, EXPN population


Population detail:

Population location: U.S.A.: All of AL, AR, CA, CO, CT, DE, FL, GA, KS, KY, LA, MA, MD, ME, MO, MS, NC, NE, NH, NJ, NV, NY, OK, PA, RI, SC, TN, VA, VT and WV; those portions of AZ, NM, and TX not included in an experimental population as set forth below; and portions of IA, IN, IL, ND, OH, OR, SD, UT, and WA as follows: (1) Southern IA, (that portion south of the centerline of Highway 80); (2) Most of IN (that portion south of the centerline of Highway 80); (3) Most of IL (that portion south of the centerline of Highway 80); (4) Western ND (that portion south and west of the Missouri River upstream to Lake Sakakawea and west of the centerline of Highway 83 from Lake Sakakawea to the Canadian border); (5) Most of OH (that portion south of the centerline of Highway 80 and east of the Maumee River at Toledo); (6) Western OR (that portion of OR west of the centerline of Highway 395 and Highway 78 north of Burns Junction and that portion of OR west of the centerline of Highway 95 south of Burns Junction); (7) Western SD (that portion south and west of the Missouri River); (8) Most of Utah (that portion of UT south and west of the centerline of Highway 84 and that portion of UT south of Highway 80 from Echo to the UT / WY Stateline); and (9) Western WA (that portion of WA west of the centerline of Highway 97 and Highway 17 north of Mesa and that portion of WA west of the centerline of Highway 395 south of Mesa). Mexico.
Listing status: E

Population location: U.S.A. (portions of AZ, NM and TX - see section 17.84(k))
Listing status: EXPN

For most current information and documents related to the conservation status and management of Canis lupus , see its USFWS Species Profile

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"Few animals have ever haunted our dreams or fired our imaginations more than the wolf. Unfortunately, by the early part of this century, man had almost exterminated the wolf from the lower 48 states. The recovery of the wolf is becoming an impressive conservation success story and a gift to future generations" (Bruce Babbitt, Secretary of the Interior).

Wolves play an important role in the ecosystem by controlling natural prey populations and removing weak individuals. Unfortunately, people who keep livestock believed that gray wolves posed a terrible threat to their herds and wolf extermination programs became widespread. Populations of gray wolves were nearly eradicated. Currently in the lower 48 United States, about 2,600 gray wolves exist, with nearly 2,000 in Minnesota (compared to the few hundred living there in the mid-20th century). Successful recovery plans have been developed throughout the country These plans evaluate populations to determine distribution, abundance, and status.  The main cause of population declines has been habitat destruction and persecution by humans. But the reintroduction of gray wolves into protected lands has greatly increased the likelihood of their survival in North America.  Populations in Alaska and Canada have remained steady and are fairly numerous. Currently the State of Alaska manages 6,000 to 8,000 gray wolves and Canada's populations are estimated at about 50,000. The wolves in Canada are managed by provincial governments and are not currently threatened.

In western Eurasia gray wolf populations have been reduced to isolated remnants in Poland, Scandinavia, Russia, Portugal, Spain, and Italy. Wolves were exterminated from the British Isles in the 1700's and nearly disappeared from Japan and Greenland in the 20th century. Greenland's wolf populations seem to have made a full recovery. The status of wolf populations throughout much of eastern Eurasia is poorly known, but in many areas populations are probably stable.

Gray wolves are listed were until recently listed as endangered by the U.S. Fish and Wildlife Service and as threatened by the state of Michigan DNR. Most U.S. populations of gray wolves have now been delisted, except for experimental populations of Mexican gray wolves in the southwest. They are in CITES Appendix II, except for populations in Bhutan, India, Nepal and Pakistan, which are in Appendix I.

IUCN Red List of Threatened Species: least concern

US Federal List: endangered

CITES: appendix i; appendix ii

State of Michigan List: threatened

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More information on the state-level protection status of gray wolves in the United States
is available at NatureServe, although recent changes in status may not be included.

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U.S. Federal Legal Status

The gray wolf is delisted in the mountain-prairie and Great Lakes-Big Rivers
regions and as an Experimental Population (nonessential) in the Southwest [30].

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"Few animals have ever haunted our dreams or fired our imaginations more than the wolf. Unfortunately, by the early part of this century, man had almost exterminated the wolf from the lower 48 states. The recovery of the wolf is becoming an impressive conservation success story and a gift to future generations" (Bruce Babbitt, Secretary of the Interior).

Wolves play an important role in the ecosystem by controlling natural prey populations and removing weak individuals. As settlement increased, the belief that livestock was endangered by wolf populations also increased. As such, the frequency of hunting the gray wolf exploded. The populations were nearly eradicated.  Currently in the lower 48 United States, about 2,600 gray wolves exist, with nearly 2,000 in Minnesota (compared to the few hundred living there in the mid-20th century). Successful recovery plans have been developed throughout the country. These plans evaluate the populations to determine distribution, abundance, and status.  The main cause of population declines has been habitat destruction and persecution by humans. But the reintroduction of gray wolves into protected lands has greatly increased the likelihood of their survival in North America.  Populations in Alaska and Canada have remained steady and are fairly numerous. Currently the State of Alaska manages 6,000 to 8,000 gray wolves and Canada's populations are estimated at about 50,000. The wolves in Canada are managed by provincial governments and are not currently threatened.

In western Eurasia gray wolf populations have been reduced to isolated remnants in Poland, Scandinavia, Russia, Portugal, Spain, and Italy. Wolves were exterminated from the British Isles in the 1700's and nearly disappeared from Japan and Greenland in the 20th century. Greenland's wolf populations seem to have made a full recovery. The status of wolf populations throughout much of eastern Eurasia is poorly known, but in many areas populations are probably stable.

Gray wolves are listed were until recently listed as endangered by the U.S. Fish and Wildlife Service and as threatened by the state of Michigan DNR. Most U.S. populations of gray wolves have now been delisted, except for experimental populations of Mexican gray wolves in the southwest. They are in CITES Appendix II, except for populations in Bhutan, India, Nepal and Pakistan, which are in Appendix I.

US Federal List: endangered

CITES: appendix i; appendix ii

State of Michigan List: threatened

IUCN Red List of Threatened Species: least concern

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Status in Egypt

Native, resident?

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Population

Population
Because of the diversity in climate, topography, vegetation, human settlement and development of wolf range, wolf populations in various parts of the original range vary from extinct to relatively pristine. Wolf densities vary from about one/12 km² to one/120 km².

Sillero et al. (2004) provide details, for each range country, on subspecies present, population status, approximate numbers, the percentage of former range occupied at present, main prey (where known), legal status, and cause of decline.

Population Trend
Stable
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Threats

Degree of Threat: B : Moderately threatened throughout its range, communities provide natural resources that when exploited alter the composition and structure of the community over the long-term, but are apparently recoverable

Comments: Exterminated from large areas through trapping, shooting, poisoning, and reduction in prey populations (ungulate herds). Threatened by direct human-caused mortality and possibly habitat loss. Landscape change resulting from development may interfere with restoration in some areas (Carroll et al. 2003). Heavily persecuted in former Soviet Union and in China.

The threats to the northern Rocky Mountain wolf population have been reduced or eliminated as evidenced by the population exceeding the numerical, distributional, and temporal recovery goals each year since 2002 (USFWS 2006).

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Major Threats
Their original worldwide range has been reduced by about one-third, primarily in developed areas of Europe, Asia, Mexico, and the United States by poisoning and deliberate persecution due to depredation on livestock. Since about 1970, legal protection, land-use changes, and rural human population shifts to cities have arrested wolf population declines and fostered natural recolonization in parts of Western Europe and the United States, and reintroduction in the western United States. Continued threats include competition with humans for livestock, especially in developing countries, exaggerated concern by the public concerning the threat and danger of wolves, and fragmentation of habitat, with resulting areas becoming too small for populations with long-term viability. There is sustainable utilization of the species' fur in Canada, Alaska, and the former Soviet Union and Mongolia.
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Management

Restoration Potential: Natural recolonization from populations to the north may not be sufficient for restoration of wolf populations in the southern Rocky Mountains. Due to predicted reductions in carrying capacity resulting from development trends and likely landscape change, active reintroduction to two sites within the southern Rocky Mountain region may be necessary to ensure low extinction probability (Carroll et al. (2003).

Management Requirements: See Mladenoff et al. (1997) for a discussion of management related to wolf recovery in the Great Lakes region versus other aspects of biodiversity (e.g., wolves depend on high densities of deer, which can have negative impacts on natural communities and their constituent species). See Cohn (1990) and Matthews and Moseley (1990) for brief discussions of issues related to reintroduction (e.g., compensation for livestock losses, wolf protection status, state vs. federal jursidiction over wolf management). See Fritts et al. (1992) for information on management of wolf-livestock conflicts in Minnesota. See USFWS (1994, Final EIS) and Federal Register (16 August 1994) for an extensive discussion of all issues related to reintroduction in Yellowstone and central Idaho. See Jhala and Giles (1991) for management recommendations for populations in northwestern India. See Theberge (1991) for information on management in Canada.

Limit access of humans and livestock in protected areas.

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Global Protection: Few to several (1-12) occurrences appropriately protected and managed

Comments: Packs and habitat are protected in at least Isle Royal National Park (Michigan), Denali National Park (Alaska), and Algonquin Provincial Park (Ontario).

Needs: Need to protect several large tracts of lands as rangeland for wolf packs.

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Conservation Actions

Conservation Actions
The species is included in CITES Appendix II, except populations from Bhutan, India, Nepal and Pakistan, which are listed on Appendix I. There is extensive legal protection in many European countries; however, enforcement is variable and often non-existent.

The species now receives legal protection in Afghanistan, having been listed by the Afghan Government on the country's 2009 Protected Species List. This prohibits all hunting and trade of C. lupus within Afghanistan.

It occurs in many protected areas across its range.

Occurrence in captivity:
The species lives and breeds well in captivity and is common in many zoological gardens.

Gaps in knowledge:
One of the most important questions still remaining about wolves involves the nature of their interaction with prey populations. The conditions under which wolves limit, regulate, or control their population is still open and important (Mech and Boitani 2003). Of more academic interest are questions involving wolf genetics, scent-marking behaviour, pseudo pregnancy and diseases (Mech 1995).
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Use of Fire in Population Management

More info for the term: cover

Fire can be used to create browse for ungulates which, in turn, provides
prey for gray wolves.  In Minnesota Heinselman [15] concluded that enough
early postfire plant communities must exist within a gray wolf pack's
territory to support a surplus of deer, moose, and beaver for prey.
Adequate hiding cover should be maintained for the ungulates.  If they
are abundant then gray wolf populations have a better chance of thriving.
Gray wolves prosper best when they have a large area, relatively free from
human disturbance, in which to roam, and when there is a surplus of
ungulates [16].  Frequent fires that promote ungulate browse in and
around areas that are at least moderately remote offer ideal gray wolf
habitat.
  • 15. Heinselman, Miron L. 1973. Fire in the virgin forests of the Boundary Waters Canoe Area, Minnesota. Quaternary Research. 3: 329-382. [282]
  • 16. Herman, Margaret, Willard, E. Earl. 1978. Rocky Mountain wolf and its habitat. Missoula, MT: U.S. Department of Agriculture, Forest Service, National Forest System Cooperative Forestry, Forestry Research, Region 1. 17 p. [16522]

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Management Considerations

Organized efforts to kill all the remaining gray wolves in the western United
States began in the 1860's.  Yellowstone and Glacier National Parks
established an official predator-control policy between 1914 and 1926
[27].  Today both parks are included in the Northern Rocky Mountain Wolf
Recovery Plan as two areas capable of sustaining viable wolf
populations.  Bunnell and Kremsater [4] concluded that wolves need about
7,818 square miles (20,250 sq km) to maintain a viable population of 50
individuals.  Fear of livestock depredation seems to be the single most
cause of opposition to gray wolf recovery.  Also hunters worry that game will
be less available if gray wolves were to recolonize their former ranges.  In
Minnesota, northwestern Montana, central Idaho, and the Greater
Yellowstone Ecosystem, livestock owners are reimbursed for animals taken by gray wolves [27].  An economic analysis conducted by Duffield
[36] concluded that gray wolf reintroduction could possibly reduce the number
of hunting permits, but that revenues lost would not exceed revenues
gained from tourism in and around Yellowstone Park, due to the increase
in photographers, filmmakers, and others wanting to see gray wolves.
  • 4. Bunnell, Fred L.; Kremsater, Laurie L. 1990. Sustaining wildlife in managed forests. Northwest Environmental Journal. 6(2): 243-269. [12830]
  • 27. Ream, Robert R.; Mattson, Ursula I. 1982. Wolf status in the northern Rockies. In: Harrington, Fred H.; Paquet, Paul C., eds. Wolves of the world: Perspectives of behavior, ecology, and conservation. Park Ridge, NJ: Noyes Publications: 362-381. [13871]
  • 36. Duffield, J. 1992. An economic analysis of wolf recovery in Yellowstone: Park visitor attitudes and values. In: Varley, J. D.; Brewster, W. G., eds. Wolves for Yellowstone? A report to the United States Congress. Vol. 4. Research and analysis. Yellowstone National Park, WY: U.S. Department of the Interior, National Park Service: pp. 2-35 to 2-87. [29422]

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Relevance to Humans and Ecosystems

Benefits

Economic Uses

Comments: See Gasaway et al. (1983) for information on wolf-prey-human relations in interior Alaska. Wolves sometimes prey on livestock or game ungulates, especially during the wolf denning period. Wolf depredation on livestock increased in the late 1980s and reached a record high level in Minnesota by 1990, and wolf sightings in a designated "no wolf" zone dramatically increased (USFWS 1990; see also Fritts et al. 1992). In recent years the USDA Animal and Plant Health Inspection Service has killed 6-42 wolves annually in livestock-depredation control actions in the northcentral U.S. (USFWS 1990). As of 1990, wolf depredation control was to continue, though possibly limited by funding shortfalls. See Fritts (1982) for information on control of predation on livestock in Minnesota. In Canada, recent annual kill by humans was estimated at 7% of population (Theberge 1991).

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Economic Importance for Humans: Negative

Gray wolves may sometimes kill livestock. The extent of livestock loss to wolves is often overstated, wolves typically prefer their wild prey.

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Economic Importance for Humans: Positive

Historically, the fur of grey wolves was used for warmth. As top predators in many ecosystems, wolves are important in controlling populations of their prey.

Wolves are important in our culture, many people believe they symbolize the spirit of wilderness. Wolf products, including posters, books, and t-shirts are very popular. Wolf ecotourism is a major source of revenue for parks and reserves.

Positive Impacts: body parts are source of valuable material; ecotourism

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Economic Importance for Humans: Negative

Gray wolves may sometimes kill livestock. The extent of livestock loss to wolves is often overstated, wolves typically prefer their wild prey.

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Economic Importance for Humans: Positive

Historically, the fur of grey wolves was used for warmth. As top predators in many ecosystems, wolves are important in controlling populations of their prey.

Wolves are important in our culture, many people believe they symbolize the spirit of wilderness. Wolf products, including posters, books, and t-shirts are very popular. Wolf ecotourism is a major source of revenue for parks and reserves.

Positive Impacts: body parts are source of valuable material; ecotourism

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Wikipedia

Dingo

For other uses, see Dingo (disambiguation).

The dingo (Canis lupus dingo) is a free-ranging dog found mainly in Australia, as well as Southeast Asia, where it is said to have originated. It is currently classified as a subspecies of the grey wolf, Canis lupus.

The dingo's habitat ranges from deserts to grasslands and the edges of forests. Dingoes will normally make their dens in deserted rabbit holes and hollow logs close to an essential supply of water.

The dingo is the largest terrestrial predator in Australia, and plays an important role as an apex predator. However, the dingo is seen as a pest by livestock farmers due to attacks on animals. Conversely, their predation on rabbits, kangaroos and rats may be of benefit to graziers.

For many Australians, the dingo is a cultural icon. There is fear of the subspecies becoming extinct. The dingo is seen by many as being responsible for thylacine extinction on the Australian mainland about two thousand years ago,[3] although a recent study challenges this view.[4] Dingoes have a prominent role in the culture of Aboriginal Australians as a feature of stories and ceremonies, and they are depicted on rock carvings and cave paintings.[5]

Nomenclature[edit]

The dingo has several names in both scientific and non-scientific literature, with "dingo" being the most commonly used. In Australia, the term "wild dog" is also widely used, but generally includes dingoes as well as dingo-hybrids and other feral dogs.[6]

Scientific name[edit]

Since its first official nomenclature in 1792 (Canis antarcticus), the scientific name of the dingo has changed several times.[7]

Current taxonomy classifies the Australian dingo, together with its closest relatives outside of Australia, as Canis lupus dingo, a subspecies of grey wolf separate from the familiar common dog, Canis lupus familiaris, while still united with familiaris as an intrataxonomic clade called "domestic dog".[7] An older taxonomy, used throughout most of the 20th century, applied the epithet Canis familiaris dingo. This taxonomy assumed that domestic dogs are a distinct species from the grey wolf, with the dingo classified as a subspecies of domestic dog. However, the term Canis dingo,[8][9] which classifies the dingo as a separate species from both dogs and wolves, has gained support in 2014 in a study that established a reference description of the dingo based on pre-20th century specimens that are unlikely to have been influenced by hybridisation.[10][11] The dingo differs from the domestic dog by relatively larger palatal width, relatively longer rostrum, relatively shorter skull height and relatively wider top ridge of skull.[10] A sample of 19th century dingo skins the study examined suggests that there was considerable variability in the colour of dingoes and included various combinations of yellow, white, ginger and darker variations from tan to black.[10] Although it remained difficult to provide consistent and clear diagnostic features, the study placed morphological limits on what can be considered a dingo.[10]

Colloquial and indigenous names[edit]

The most commonly used name is dingo, which has its origins in the early European colonisation in New South Wales and is most likely derived from the word tingo, used by the Aboriginal people of Port Jackson for their camp dogs.[12] Depending on where they live, local dingoes can be called "alpine dingoes," "desert dingoes," "northern dingoes," "Cape York dingoes," or "tropical dingoes". More recently, people have begun to call dingoes "Australian native dogs" or, by reasoning that they are a subspecies of Canis lupus, "Australian wolves".[13]

The dingo has been given different names in the Indigenous Australian languages, including joogong, mirigung, noggum, boolomo, papa-inura, wantibirri, maliki, kal, dwer-da, kurpany, aringka, palangamwari, repeti and warrigal.[6] Some languages have different names for the dingoes depending on where they live; the Yarralin, for instance, call the dingoes that live with them walaku and those in the wilderness ngurakin.[5]

Description[edit]

The skeleton of a dingo at the Royal Veterinary College

Domestic and pariah dogs in southern Asia share so many characteristics with Australian dingoes that they are now considered to be members of the same taxon Canis lupus dingo, a particular subspecies of Canis lupus. While the relationship with humans varies widely among these animals, they are all quite similar in terms of physical features.[6]

Dingo skull illustrated by Frédéric Cuvier: As is typical in domestic dogs, the dingo's relative brain size is smaller than that of all non-domesticated subspecies of wolves, being almost identical in size to that of dogs considered Canis lupus familiaris.[14]

A dingo has a relatively broad head, a pointed muzzle and erect ears. Eye colour varies from yellow over orange to brown.[15] Compared to other similarly sized familiaris dogs, dingoes have longer muzzles, larger carnassials (large teeth found in many carnivorous mammals), longer canine teeth, and flatter skulls with larger nuchal lines.[12]

Size[edit]

The average Australian dingo is 52 to 60 cm (20 to 24 in) tall at the shoulders and measures 117 to 154 cm (46 to 61 in) from nose to tail tip. The average weight is 13 to 20 kg (29 to 44 lb); however, there are a few records of outsized dingoes weighing up to 27 to 35 kg (60 to 77 lb).[16][17] Males are typically larger and heavier than females of the same age. Dingoes from northern and northwestern Australia are larger than central and southern populations. Australian dingoes are invariably heavier than Asian ones.[6] The legs are about half the length of the body and the head put together. The hind feet make up a third of the hind legs and have no dewclaws.[6] Dingoes can have sabre-form tails (typically carried erect with a curve towards the back) or tails carried directly on the back.[15]

Fur[edit]

Typical dingo colouration

Fur of an adult dingo is short and soft, bushy on the tail, and varies in thickness and length depending on the climate. The fur colour is mostly sandy to reddish brown, but can include tan patterns and sometimes be black, light brown, or white. Completely black dingoes might have been more prevalent in Australia in the past, but have only been rarely sighted in recent times. They are now more common in Asia.[12]

Most dingoes are at least bi-coloured, with small, white markings on the chest, muzzle, tag, legs and paws being the most common feature. Pure Dingoes are also found in white or cream (not Albinism) they are also found in Black & Tan colourations. In the case of reddish individuals, there can be small, distinctive, dark stripes on the shoulders.[6]

Origin and genetic status[edit]

Since dingoes were the largest wild placental mammals in Australia at the time of colonisation and looked similar to domestic dogs, their origin has always been questioned and much debated. Archaeological and morphological studies indicated a relatively late introduction and a close relationship to other domestic dogs. Their exact descent, place of origin and date of arrival in Australia were not identified, nor whether they had once been domesticated or half-domesticated and had gone feral, or whether they had already existed as truly wild animals.[18]

It is widely held that dingoes have evolved or were bred from the Indian wolf or Arabian wolf around 6,000 to 10,000 years ago, as was assumed for all domestic dogs.[19] This theory was based on the morphological similarities of dingo skulls and the skulls of these subspecies of wolves. However, genetic analyses indicated a much earlier domestication.

DNA analysis[edit]

Analyses of amino acid sequences of the haemoglobin from a "pure" dingo in the 1970s supported the theory that dingoes are more closely related to domestic dogs than they are to grey wolves or coyotes. As a result, it was assumed that dingoes and other similar Asian dogs belong to a group of domestic dogs that went feral at a very early time. DNA studies on Australian dingoes and domestic dogs were also undertaken in an effort to reliably differentiate between both populations and to try to determine the extent of interbreeding.

The first two examinations looked firstly at 14 loci (the specific locations of the DNA-sequence of a chromosome), with five of these being more closely examined. No genetic difference could be found. The analyses were then extended to cover 16 loci, comparing dingoes from Central Australia, dingoes from the Eastern Highlands, dingo-hybrids and domestic dogs of other origin. Again, no differences could be found, regardless of the type of examination used. It was reasoned that dingoes and domestic dogs must have a very similar gene pool. However, since only a few differences in the enzymes of different species of the genus Canis could be found, it was assumed that a lack of differences might not indicate a close taxonomic relationship. It was also reasoned that the degree of interbreeding in the wild would be hard to determine.[20]

During further analyses in the late 1990s, researchers examined 14 loci and detected a significantly lower genetic variability among Australian dingoes than among domestic dogs, leading to consideration of the possibility of a small founding population. There was one locus found that might have been suitable for differentiation, but not in the case of interbreeding of a dingo-hybrid with a "pure" dingo. Additionally, it was suspected that findings of other suitable loci might be used to determine whether there are clearly separate sub-populations of the "pure" dingoes.[21]

Mitochondrial DNA sequences[edit]

To determine the origin and time of arrival of Australian dingoes, mitochondrial DNA (mtDNA) sequences of 211 dingoes and 19 archaeological samples from pre-European Polynesia were compared with mtDNA samples of 676 domestic dogs and 38 grey wolves in 2004. The domestic dog samples came from China, Africa, Southwest Asia, India, Siberia, the arctic America, Europe, Mongolia, Korea, Japan, Vietnam, Cambodia, Thailand, Indonesia, the Philippines, Malaysia, New Zealand, Hawaii and the highlands of New Guinea. The dingo samples came from zoos, wildlife parks, dingo conservation groups, dingo lovers and 192 wild living specimens from 27 areas in Australia, mainly located in the Pilbara region, New South Wales and northeastern Victoria. The wild specimens had been selected based on similarities of external appearance to exclude the influence of dingo-hybrids and domestic dogs as far as possible.

Compared to wolves and domestic dogs, the variation of mtDNA sequences was very limited. Among dingoes, only 20 mtDNA sequences differing in two point mutations at most could be found. In comparison, 114 mtDNA-sequences with a maximal difference of 16 point mutations between the DNA-types could be found among domestic dogs. Two of the dingo mtDNA-types were similar to that of domestic dogs (A9, A29), while the other 18 types were unique to dingoes.

In a phylogenetic tree of wolves and domestic dogs, dingoes were included in the main clade (A), which contained 70% of all domestic dog types. Within this clade, the dingo-types formed a group around the type A29, which was surrounded by 12 less frequent dingo-types, as well as a set of other domestic dog types. This mtDNA-type was found in 53% of the dingoes and was also found among some domestic dogs from East Asia, New Guinea and the American Arctic. Based on these findings, it was reasoned that all dingo-mtDNA-types originated in A29. A9 was only found in one individual, and it was regarded as possible that this type is the result of a parallel mutation.

Based on a mutation rate of mtDNA with A29 being the only founder type, it was considered that dingoes probably arrived in Australia between 4,600 and 5,400 years ago, which was consistent with archaeological findings. However, it was also considered that dingoes might have arrived from an even earlier date of up to 10,800 years ago in the event of the mtDNA-mutation rate being slower than assumed. It was further reasoned that these findings strongly indicate a descent of dingoes from East Asian domestic dogs and not from Indian domestic dogs or from wolves. In addition these findings indicated two possibilities of descent: all Australian dingoes are descended from a few domestic dogs, theoretically one pregnant female; and all Australian dingoes are descended from a group of domestic dogs, who radically lost their genetic diversity through one or several severe genetic bottlenecks on their way from the Asian continent over Southeast Asia.

Other types[edit]

Nonetheless, the existence of other mtDNA-types on the islands surrounding Australia indicate there have been other types apart from A29 and only one single founding event. These results also indicated that there hasn't been any significant introduction of other domestic dog on the Australian continent prior to the arrival of the Europeans. Also, a shared origin and some sort of genetic exchange between Australian dingoes and the New Guinea singing dogs was regarded as possible. The current state of the Australian dingoes was ascribed to the long wild existence of these dogs and assumed that they are an isolated example of early domestic dogs.

Despite accordant claims,[22][23] these findings did not show that only dingo females mate with non-dingo males and not vice versa. The findings would not allow such a conclusion, since the mating of a dingo female with a non-dingo male could not be detected via analyses of mtDNA. Furthermore the researchers made sure from the start that dingo-hybrids were excluded as far as possible.[18]

Communication[edit]

Like all domestic dogs, dingoes tend towards phonetic communication. However, in contrast to domestic dogs, dingoes howl and whimper more, and bark less. Eight sound classes with 19 sound types have been identified.[24]

Barking[edit]

Compared to most domestic dogs, the bark of a dingo is short and monosyllabic, and is rarely used. Barking was observed to make up only 5% of vocalisations. Dog barking has always been distinct from wolf barking.[25] Australian dingoes bark mainly in swooshing noises or in a mixture of atonal and tonal sounds. In addition, barking is almost exclusively used for giving warnings. Warn-barking in a homotypical sequence and a kind of "warn-howling" in a heterotypical sequence have also been observed. The bark-howling starts with several barks and then fades into a rising and ebbing howl and is probably (similar to coughing) used to warn the puppies and members of the pack. Additionally, dingoes emit a sort of "wailing" sound, which they mostly use when approaching a wateringhole, probably to warn already present dingoes.[6]

According to the present state of knowledge, it is not possible to get Australian dingoes to bark more frequently by putting them in contact with other domestic dogs. However, German zoologist Alfred Brehm reported a dingo that learned the more "typical" form of barking and how to use it, while its brother did not.[26] Whether dingoes bark or bark-howl less frequently in general is not certain.[24]

Howling[edit]

Captive dingoes howling

Dingoes have three basic forms of howling (moans, bark-howls and snuffs) with at least 10 variations. Usually, three kinds of howls are distinguished: long and persistent, rising and ebbing, and short and abrupt.

Observations have shown that each kind of howling has several variations, though their purpose is unknown. The frequency of howling varies with the season and time of day, and is also influenced by breeding, migration, lactation, social stability and dispersal behaviour. Howling can be more frequent in times of food shortage, because the dogs become more widely distributed within their home range.[6]

Additionally, howling seems to have a group function, and is sometimes an expression of joy (for example, greeting-howls). Overall howling was observed less frequently in dingoes than among grey wolves. It may happen that one dog will begin to howl, and several or all other dogs will howl back and bark from time to time. In the wilderness, dingoes howl over long distances to attract other members of the pack, to find other dogs, or to keep intruders at bay. Dingoes howl in chorus with significant pitches, and with increasing number of pack-members, the variability of pitches also increases.[27] Therefore, it is suspected that dingoes can measure the size of a pack without visual contact.[12] Moreover, it has been proposed that their highly variable chorus howls may generate a confounding effect in the receivers by making pack size appear larger.[28]

Other forms of communication[edit]

Growling, making up approximately 65% of the vocalisations, is used in an agonistic context for dominance, and as a defensive sound. Similar to many domestic dogs, a reactive usage of defensive growling is only rarely observed. Growling very often occurs in combination with other sounds, and has been observed almost exclusively in swooshing noises (similar to barking).[24]

During observations in Germany, dingoes were heard to produce a sound that observers have called Schrappen. It was only observed in an agonistic context, mostly as a defence against obtrusive cubs or for defending resources. It was described as a bite intention, during which the receiver is never touched or hurt. Only a clashing of the teeth could be heard.[24]

Aside from vocal communication, dingoes communicate, like all domestic dogs, via scent marking specific objects (for example, Spinifex) or places (such as waters, trails and hunting grounds) using chemical signals from their urine, feces and scent glands. Males scent-mark more frequently than females, especially during the mating season. They also scent-rub, whereby a dog rolls its neck, shoulders, or back on something that is usually associated with food or the scent markings of other dogs.[6]

Unlike wolves, dingoes can react to social cues and gestures from humans.[29]

Behaviour[edit]

Dingoes tend to be nocturnal in warmer regions, but less so in cooler areas. Their main period of activity is around dusk and dawn. The periods of activity are short (often less than one hour) with short times of resting. Dingoes have two kinds of movement: a searching movement (apparently associated with hunting) and an exploratory movement (probably for contact and communication with other dogs).[30][31]

In general, dingoes are shy towards humans. However, there are reports of dingoes that were agitated by the presence of humans, such as around camps in national parks, near streets or suburbs.[16][32] According to studies in Queensland, the wild dogs (dingo hybrids) there, move freely at night through urban areas and cross streets and seem to get along quite well.[33]

Dietary habits[edit]

About 170 species (from insects to buffalo) have been identified as part of the dingo's diet. In general, livestock seems to make up only a small proportion of their diet.[6] In continent-wide examinations, 80% of the diet of wild dogs consisted of 10 species: red kangaroo, swamp wallaby, cattle, dusky rat, magpie goose, common brushtail possum, long-haired rat, agile wallaby, European rabbit and the common wombat. This narrow range of major prey indicates these wild dogs are rather specialised,[12] but in the tropical rainforests of northeastern Australia, dingoes are supposed to be opportunistic hunters of a wide range of mammals.[34] In certain areas, they tend to specialise on the most common prey, with a preference for medium- to large-sized mammals. Their consumption of domestic cats has also been proven.[35] Non-mammalian prey is irregularly eaten and makes up only 10% of the dingo's diet. Big reptiles are only rarely captured, at least in eastern Australia, although they are widespread. It is possible that especially large monitor lizards are too defensive and well-armed, or they are simply able to flee fast enough into dens or climb trees.[12]

A dingo near the Dingo Fence

Dietary composition varies from region to region. In the gulf region of Queensland, feral pigs and agile wallabies are the dingo's main prey. In the rainforests of the north, the main prey consists of magpie geese, rodents and agile wallabies. In the southern regions of the Northern Territory, the dogs mainly eat European rabbits, rodents, lizards and red kangaroo; in arid Central Australia, rabbits, rodents, lizards, red kangaroo and cattle carcasses; and in the dry northwest, eastern wallaroos and red kangaroo. In the deserts of the southwest, they primarily eat rabbits, and in the eastern and southeastern highlands, they eat wallabies, possums and wombats.

To what extent the availability of rabbits influences the composition of the diet cannot be clarified. However, because rabbit haemorrhagic disease killed a large part of the Australian rabbit population at the end of the 20th century, it is suspected that the primary prey of the dogs has changed in the affected areas. Also, on Fraser Island, fish have been proven to be a part of the dingo diet. The main prey species, though, are bandicoots and several rodents. Dingoes also eat a lot of echidnas, crabs, small skinks, fruits and other plants, as well as insects (mostly beetles). During these observations, only 10% of the examined feces-samples contained human garbage (in earlier studies 50% were reported).[12]

When scavenging for food, wild dogs (presumably, all dogs free to roam, not just dingoes) primarily eat cattle and kangaroo carcasses. Dingoes in coastal regions regularly patrol the coast for dead fish, seals, penguins and other washed-up birds.[12]

Dingoes in general drink one litre of water a day in the summer and about half a litre a day in winter. During the winter in arid regions, dingoes could potentially live from the liquid in the bodies of their prey, as long as the number of prey is sufficient. Similarly, weaned cubs in Central Australia are able to draw their necessary requirements of liquid from their food. There, regurgitation of water by the females for the cubs was observed. During lactation, females have no higher need of water than usual, since they consume the urine and feces of the cubs and therefore recycle the water and keep the den clean.[12]

Hunting behaviour[edit]

Dingoes often kill by biting the throat, and they adjust their hunting strategies to suit circumstances. For larger prey, due to strength and potential danger, two or more individuals are needed to bring down the prey. Such group formations are unnecessary when hunting rabbits or other small prey.[12]

Kangaroo hunts are probably more successful in open areas than in places with high densities of vegetation, and juvenile kangaroos are killed more often than adults. Dingoes typically hunt large kangaroos by having lead dingoes chase the quarry toward their waiting packmates, which are skilled at cutting corners in chases. In one area of Central Australia, dingoes hunt kangaroos by chasing them toward a wire fence that hindered their escape.[27]

Birds can be captured when they do not fly or fail to take off fast enough. Dingoes also steal the prey of eagles and the coordinated attack of three dingoes for killing a large monitor lizard has been observed.[36]

Reports state that some dingoes live almost entirely on human food through stealing, scavenging, or begging. In fact, dingoes are well known for such behaviour in some parts of Australia. It is suspected that this might cause the loss of hunting strategies or a change in the social structures.[37]

During studies at the Fortescue River in the mid-1970s, observation showed that most of the studied dingoes learned to hunt and kill sheep very quickly, even without prior contact with sheep. Although the dingoes killed many sheep at that time, they still killed and ate kangaroos.

During the early 1990s, wild dogs were observed to have an extraordinarily high success rate when killing sheep, and did not have to hunt in a coordinated manner to achieve success. Often, a dog may chase and outrun a single sheep, only to turn away suddenly and chase another. Therefore, only a small proportion of the injured or killed sheep and goats are eaten, which seems to be the rule and not the exception. The dog probably falls into some kind of "killing spree," due to the rather panicked and uncontrolled flight behaviour of the sheep, which run in front of the dingoes time and again and, therefore, cause one attack after another. Dingoes often attack sheep from behind during the sheep's flight, which causes injuries to the sheep's hind legs. Rams are normally attacked from the side – probably in order to avoid the horns – or sometimes on the testicles. Inexperienced dingoes, or those that kill "for fun," sometimes cause significant damage to the sheep's hind legs, which often causes death.[38][39]

Nearly all dingo attacks on cattle and water buffalo are directed against calves. Hunting success depends on the health and condition of the adult bovines and on their ability to defend their calves. The defence behaviour of the mother can be sufficient to fend off an attack. Therefore, the basic dingo tactics of attack are distracting the mother, rousing the herd/group and waiting (sometimes for hours), and testing of the herd to find the weakest members.

While locating a cattle herd, dingoes have been observed to make several feint attacks, during which they concentrate on the calves at first then, later on, attack the mothers to distract them. Thereupon, the dingoes retreat and wait at a distance from the herd until the rest of the cows have gathered their calves and move on.

During another observed attack, "subgroups" of a dingo pack took turns in attacking and resting, until the mother was too tired to effectively defend her calf. Dingoes have been observed hunting a water buffalo with an estimated weight of 200 kg, and taking turns biting the buffalo's legs during the chase.[12][40]

Social behaviour[edit]

The dingo's social behaviour is about as flexible as that of a coyote or gray wolf, which is perhaps one of the reasons it was initially believed that the dingo was descended from the Indian wolf.[41] While young males are often solitary and nomadic in nature, breeding adults will often form a settled pack.[42] However, in areas of the dingo's habitat with a widely spaced population, breeding pairs remain together, apart from others.[42]

Where conditions are favourable among dingo packs, the pack is stable with a distinct territory and little overlap between neighbors.[43] The size of packs often appears to correspond to the size of prey that appears in the pack's territory.[43] Desert areas have smaller groups of dingoes with a more loose territorial behaviour and sharing of the water sites.[44] It has been noted that the average monthly pack size was between three and twelve members.[45]

Similar to other canids, a dingo pack largely consists of a mated pair, their current year's offspring, and occasionally a previous year's offspring.[43] There are dominance hierarchies both between and within males and females, with males usually being more dominant than females.[43] A few exceptions have been noted in captive packs, however.[43] During travel, while eating prey, or when approaching a water source for the first time, the breeding male will be seen as the leader, or alpha.[46] Subordinate dingoes will approach a more dominant dog in a slightly crouched posture, ears flat and tail down, to ensure peace in the pack.[43] Establishment of artificial packs in captive dingoes have failed.[43]

Reproduction[edit]

Dingo pups from the Tierpark Berlin

Dingoes breed once annually, depending on the estrus cycle of the females which, according to most sources, only come in heat once per year. Dingo females can come in heat twice per year, but can only be pregnant once a year, with the second time only seeming to be pregnant.[47][48]

Males are virile throughout the year in most regions, but have a lower sperm production during the summer in most cases. During studies on dingoes from the Eastern Highlands and Central Australia in captivity, no specific breeding cycle could be observed. All were potent throughout the year. The breeding was only regulated by the heat of the females. A rise in testosterone was observed in the males during the breeding season, but this was attributed to the heat of the females and copulation. In contrast to the captive dingoes, captured dingo males from Central Australia did show evidence of a male breeding cycle. Those dingoes showed no interest in females in heat (this time other domestic dogs) outside of the mating season (January to July) and did not breed with them.[49]

The mating season usually occurs in Australia between March and May (according to other sources between April and June). In Southeast Asia, mating occurs between August and September. During this time, dingoes may actively defend their territories using vocalisations, dominance behaviour, growling and barking.[12]

Most females in the wild start breeding at the age of two years. Within packs, the alpha female tends to go into heat before subordinates and actively suppresses mating attempts by other females. Males become sexually mature between the ages of one to three years. The precise start of breeding varies depending on age, social status, geographic range and seasonal conditions. Among dingoes in captivity, the pre-estrus was observed to last 10–12 days. However, it is suspected that the pre-estrus may last as long as 60 days in the wild.[6]

A male dingo with his pups

In general, the only dingoes in a pack that successfully breed are the alpha pair, and the other pack members help with raising the pups. Subordinates are actively prevented from breeding by the alpha pair and some subordinate females have a false pregnancy. Low-ranking or solitary dingoes can successfully breed if the pack structure breaks up.[50]

The gestation period lasts for 61–69 days and the size of the litter can range from one to 10 (usually five) cubs, with the number of males born tending to be higher than that of females. Pups of subordinate females usually get killed by the alpha female, which causes the population increase to be low even in good times. This behaviour possibly developed as an adaptation to the fluctuating environmental conditions in Australia. Pups are usually born between May and August (the winter period), but in tropical regions, breeding can occur at any time of the year.[6]

At the age of three weeks, the pups leave the den for the first time, and leave it completely at eight weeks. In Australia, dens are mostly underground. There are reports of dens in abandoned rabbit burrows, rock formations, under boulders in dry creeks, under large spinifex, in hollow logs, in augmented burrows of monitor lizards and wombat burrows. The pups usually stray around the den within a radius of 3 km, and are accompanied by older dogs during longer travels. The transition to consuming solid food is normally accompanied by all members of the pack during the age of 9 to 12 weeks. Apart from their own experiences, pups also learn through observation.[51] Young dingoes usually become independent at the age of three to six months or they disperse at the age of 10 months when the next mating season starts.

Migration[edit]

Dingoes usually remain in one area and do not undergo seasonal migrations. However, during times of famine, even in normally "safe" areas, dingoes travel into pastoral areas, where intensive, human-induced control measures are undertaken. It was noted in Western Australia in the 1970s that young dogs can travel for long distances when necessary. About 10% of the dogs captured—all younger than 12 months—were later recaptured far away from their first location. Among these, 10% of the travelled distance for males was 21.7 km and for females 11 km.[52] Therefore, travelling dingoes had lower chances of survival in foreign territories, and it was apparently unlikely that they would survive long migrations through occupied territories. The rarity of long migration routes seemed to confirm this. During investigations in the Nullarbor Plain, even longer migration routes were recorded. The longest recorded migration route of a radio-collared dingo was about 250 km.

Mortality and health[edit]

Documented evidence shows that dingoes in captivity have survived for up to 24 years.[15]

The main cause of death for dingoes is being killed by humans, crocodiles and dogs, including other dingoes. Other causes of death include starvation and dehydration during times of drought or after strong bush fires, infanticide, snake bites, killing of cubs by wedge-tailed eagles, and injuries caused by cattle and buffalo.

Dingoes are susceptible to the same diseases as domestic dogs. At present, 38 species of parasites and pathogens have been detected in Australian dingoes. The bulk of these diseases have a minimal influence on their survival. The exceptions include canine distemper, hookworms and heart worms in North Australia and southeastern Queensland. Dingo pups can also be killed by lungworms, whipworms, hepatitis, coccidiosis, lice and ticks. Sarcoptic mange is a widespread parasitic disease among the dingoes of Australia, but is seldom debilitating. Free-roaming dogs are the primary host of Echinococcosis (tapeworms) and have an infection rate of 70 to 90%.

Distribution[edit]

It is only possible to give a crude description of the dingo's distribution area and the accordant population density. Giving an exact assessment of the distribution of dingoes and other domestic dogs is difficult since the exact extent of interbreeding between the two is not known. The following information on the distribution of the dingo applies to dogs classified as dingoes based on fur colour, body form and breeding cycle. Therefore, the maps illustrating their distribution might be conflicting.

Distribution in the past[edit]

Based on fossil, molecular and anthropogenic evidence, dingoes might have once had a widespread distribution. These early dingoes would have associated with nomadic hunter-gatherer societies and later with the rising agricultural centres. It is further assumed that they would have been tamed there and then transported to various places in the world. Findings of dingo habitation in Thailand and Vietnam, regarded as the oldest findings, have been estimated at 5,000 to 5,500 years old. The age of similar findings from the highlands of Indonesia vary from 2,500 to 5,000 years.

An early European illustration of a dingo, from Arthur Phillip's Voyage to Botany Bay in 1789. The pictured specimen was a female taken alive by Governor Phillip and given to the Marquess of Salisbury, at Hatfield House.[53]

Originally, the dingo was suspected to have been introduced to Australia in the Pleistocene by Aborigines, which led to confusion concerning the dingo's nomenclature. Today, the most common theory is that the dingo arrived in Australia about 4,000 years ago. In 1979, an eroding dingo skeleton was excavated by Brown and Gollan (ANU) on the mid-coast of southern New South Wales,[54] dated to 6,000 years of age. More recent mitochondrial DNA research estimates the arrival of dingoes to be between 4,600 and 18,300 years.[55] Evidence of dingoes appears to be absent from Tasmania, which was separated from the main Australian landmass around 12,000 years ago due to a rise in sea level, which led to the theory that dingoes have not been in Australia longer than this time. To reach Australia from Asia, there would have been at least 50 km of open sea to be crossed, even at the lowest sea level. Since there are few, if any, cases of a large land animal making such a journey by itself (the Falkland Islands wolf being a possible exception), the ancestors of modern dingoes most likely were brought to Australia on boats by Asian seafarers.[18] A dance of the Aborigines in the coastal regions of the Kimberley, during which they depict dogs running excitedly up and down a boat and finally jumping into the water, is seen as further evidence for the introduction of dingoes by seafarers.[56] These dogs possibly were used as food or eventually guard dogs. Potentially, the dingo came to Australia and the islands of Southeast Asia and the Pacific during the course of expansion of the Austronesian culture.

Theoretical dingo migration routes

The two main theories concerning the geographical origin and travel routes of the modern dingo's ancestors and their arrival in Australia are:[57]

  • The dingoes originated in East Asia and took a travel route over the Southeast Asian islands due to their close proximity to Australia, and the relatively easy accessibility over the islands of the Southeast Asian archipelago. This theory is supported by examination of the mtDNA of Australian dingoes.[18]
  • Sheepdogs were introduced from the Indus Valley in Asia, over Timor by Indian seafarers, based on similarities in skeletal anatomy of Indian pariah dogs and Iranian wolves. This theory implies that the oldest-known fossils are 4,000 years old and were found on Timor, where the dogs coexisted for a time with pigs and sheep. This theory would be supported by the assumption that the simultaneous appearance of certain stone tools was caused by Indian influence, but other authorities dispute this. Recent genetic research on aboriginal DNA seems to support this conclusion, that Indian seafarers brought their dogs and other tools with them to Australia 4,000 years ago.[58][59][60]

Whether there were several introductions of dingoes to Australia or just one is not yet known.

The first official report of a "wild dog" in Australia comes from Captain William Dampier in 1699.[61] At the time, dingoes were probably[citation needed] widespread over the main part of the continent and lived in the wild, as well as alongside the Aboriginals. They were mostly tolerated by the European settlers and sometimes kept as pets. The number of dingoes was probably[citation needed] low in those times and increased since then in some parts of Australia. Their numbers probably[citation needed] increased strongly around the 1880s due to the establishment of the pastoral economy and artesian watering places, and probably[citation needed] peaked in the 1930s and 1950s. Afterwards, the numbers remained high, but the percentage of dingo-hybrids has significantly increased since then[citation needed].

Present distribution[edit]

Possible distribution of the dingo (red): The red area in Papua New Guinea shows the possible distribution of the Hallstrom dog.

Today, dingoes live in many diverse habitats, including the snow-covered mountain forests of eastern Australia, the deserts of Central Australia, and Northern Australia's tropical forest wetlands. The absence of dingoes in many parts of the Australian grasslands is probably due to human persecution. Based on skull characteristics, size, fur colour and breeding cycles, distinct regional populations could be seen to exist between Australia and Asia, but not within Australia.[6][12]

The wild dog population of Australia now includes dingoes and a wide panoply of feral domestic dogs (mostly mixed-breeds and dingo-hybrids) having an enormous variety of colours. Due to the increased availability of water, native and introduced prey, livestock and human-provided food, this population is on the increase. Reports from some parts of Australia indicate that wild dogs now hunt in packs there, where they had previously been solitary hunters.[62] Dingo densities have been measured at up to 0.3 per square kilometre in both the Guy Fawkes River region of New South Wales and in South Australia at the height of a rabbit plague.[12]

"Pure"[63] dingoes are regarded as widespread in Northern, North West and Central Australia; rare in Southern and Northeast Australia; and possibly extinct in the Southeastern and Southwestern areas.

Distribution map of Australian dingoes: The black line represents the Dingo Fence (after Fleming et al. 2001).

The establishment of agriculture caused a significant decrease in dingo numbers, and dingoes were practically expelled from the territories occupied by the sheep industry, primarily affecting large parts of southern Queensland, New South Wales, Victoria and South Australia. This situation was maintained by the construction of the Dingo Fence. Although dingoes were eradicated from most areas south of the Dingo Fence, they still exist in an area of about 58,000 km2 in the dry northern areas north of the Dingo Fence and, therefore, on about 60% of the entire area.[clarification needed]

In Victoria, wild dog populations are currently concentrated on the densely forested areas of the Eastern Highlands, from the border to New South Wales, south to Healesville and Gembrook. They also exist in the large desert in the northwest of the state. Wild dog populations in New South Wales primarily exist along the Great Dividing Range and the hinterlands on the coast, as well as in the Sturt National Park in the northwest of the state.

In the rest of the continent, dingoes are regarded as widespread, with the exception of the arid eastern half of Western Australia. In the bordering areas of South Australia and the Northern Territory, they are regarded as naturally scarce. Wild dogs are widespread in the Northern Territory, with the exception of the Tanami and Simpson Deserts, where they are rare due to the lack of watering holes. However, local concentrations exist there near artificial water sources. According to DNA examinations from 2004, the dingoes of Fraser Island are "pure".[64] However, skull measurements from the mid-1990s had a different result.[65] A 2013 study showed that dingoes living in the Tanami Desert are among the "purest" in Australia.[66]

Outside Australia, dingoes were proven to exist in Thailand, based on comparisons between the skulls of Thai dogs and those of fossil and present-day dingoes. The population there probably has the largest proportion of "pure" dingoes. They are widespread in northern and central Thailand and rare in the southern regions. They may also exist in Burma (Myanmar), China, India, Indonesia, Laos, Malaysia, Papua New Guinea, the Philippines and Vietnam, but if they exist there, their distribution is unknown. Dingoes are regarded as widespread in Sulawesi, but their distribution in the rest of Indonesia is unknown. They are regarded as rare in the Philippines and are probably extinct on many islands. In Korea, Japan and Oceania, a few local dog breeds with dingo-like features exist, but dingoes are considered extinct there.[6]

Ecological impact of the dingo after its arrival in mainland Australia[edit]

The dingo is suspected to have caused the extinction of the thylacine, the Tasmanian devil and the Tasmanian Native-hen from mainland Australia, since a correlation in space and time is found between the arrival of the dingo and the extinctions of these species. Recent studies have questioned this theory, suggesting that climate change and increasing human populations may have been the cause.[67] Dingoes do not seem to have had the same ecological impact that the red fox had in later times. This might be connected to the dingo's way of hunting and the size of their favoured prey, as well as to the low number of dingoes in the time before European colonisation.[68]

The assumption that dingoes and thylacines were competitors for the same prey stems from their external similarities; the thylacine had a stronger and more efficient bite, but was probably dependent on relatively small prey, while the dingo's stronger skull and neck would have allowed it to bring down bigger prey.[69] The dingo was probably a superior hunter, as it hunted cooperatively in packs and could better defend resources, while the thylacine was probably more solitary. Also, wild dingo populations might have had demographic support from conspecific living with humans and may have introduced new diseases that affected the thylacine more severely.[citation needed]

The extinction of the thylacine on the continent around 2,000 years ago has also been linked to changes in climate and land use by the Aborigines. It is plausible to name the dingo as the cause of the extinction, but significant morphological differences between the two suggest that the ecological overlapping of both species might be exaggerated. The dingo has the dentition of a generalist, while the thylacine had the dentition of a specialist carnivore without any signs of consumption of carrion or bones. It is also argued that the thylacine was a flexible predator that should have withstood the competition by the dingo, but was instead wiped out due to human persecution.[citation needed]

This theory does not explain how the Tasmanian devil and the dingo coexisted on the same continent until about 430 years ago, when the dingo supposedly caused the Tasmanian devil's demise. The group dynamics of dingoes should have successfully kept devils away from carrion, and since dingoes are able to break bones, little would have been left for the devils to scavenge. Additionally, devils are successful hunters of small- to medium-sized prey, so there should have been an overlapping of the species in this area, too. Furthermore, the arguments that the dingo caused the extinction of the thylacine, the devil and the hen are in direct conflict with each other. If the dingo were really so similar to the thylacine and the Tasmanian devil in its ecological role and suppressed both, then coexisting with both for such an extended time is strange. Although this is a possible result of the dingo's introduction, critics regard the evidence for this as insubstantial.[70]

Impact[edit]

Reliable information about the exact ecological, cultural and economic impact of wild dogs does not yet exist. Furthermore, the impact of wild dogs depends on several factors, and a distinction between dingoes and other domestic dogs is not necessarily made.

The appearance of a wild dog is sometimes very important when it comes to the cultural and economical impact. Here, it is often desired that the wild dog's appearance complies to what is demanded, that it is a "pure" dingo or at least looks like one.[71] In the case of their economic impact, their appearance only seems to be important when "pure" dingoes are used as a tourist attraction. Where wild dogs are regarded as pests, their appearance is only of minor importance, if at all.

The impact wild dogs have in urban areas and whether they are a danger to humans (such as direct attacks or diseases) is currently unknown.

Ecological impact[edit]

The dingo is regarded as part of the native Australian fauna by many environmentalists and biologists, as these dogs existed on the continent before the arrival of the Europeans and a mutual adaption of the dingoes and their surrounding ecosystems had occurred. However, the contrary view has dingoes as just another introduced predator that are only native to Thailand.[72]

Much of the present place of wild dogs in the Australian ecosystem, especially in the urban areas, remains unknown. Although the ecological role of dingoes in Northern and Central Australia is well understood, the same does not apply to the role of wild dogs in the east of the continent. In contrast to some claims,[73] dingoes are assumed to have a positive impact on the environment.[citation needed]

Dingoes are regarded as apex predators and possibly perform an ecological key function. It is likely (with increasing evidence from scientific research) that they control the diversity of the ecosystem by limiting the number of prey and keeping the competition in check. Wild dogs hunt feral livestock such as goats and pigs, as well as native prey and introduced animals. The low number of feral goats in Northern Australia possibly is caused by the presence of the dingoes, but whether they control the goats' numbers or not is still disputable. Studies from 1995 in the northern wet forests of Australia found the dingoes there did not reduce the number of feral pigs, but their predation only has an impact on the pig population together with the presence of water buffalos (which hinder the pigs' access to food).[74] Mike West, former president of Birds Queensland, blames dingoes for cutting down the number of Black-breasted Buttonquails on Inskip Point on Fraser Island to one.[75] West suggests that dingoes and wild dogs should be trapped; pure dingoes would be relocated and non-pure dingoes and wild dogs would be killed.[76]

Observations concerning the mutual impact of dingoes and red fox and cat populations suggest dingoes limit the access of foxes and cats to certain resources. As a result, it is assumed that a disappearance of the dingoes may cause an increase of red fox and feral cat numbers and, therefore, a higher pressure on native animals. These studies found the presence of dingoes is one of the factors that keep fox numbers in an area low, and therefore reduces pressure on native animals, which then do not disappear from the area. The countrywide numbers of red foxes are especially high where dingo numbers are low, but other factors might responsible for this, depending on the area.[35] Evidence was found for a competition between wild dogs and red foxes in the Blue Mountains of New South Wales, since there were many overlaps in the spectrum of preferred prey, but there was only evidence for local competition, not on a grand scale.[77]

It is also possible that dingoes can live with red foxes and feral cats without reducing their numbers in areas with sufficient food resources (for example, high rabbit numbers) and hiding places. Nearly nothing is known about the relationship of wild dogs and feral cats, except both mostly live in the same areas. Although wild dogs also eat cats, it is not known whether this has an impact on the cat populations.[35] At the moment, the Invasive Animals Cooperative Research Centre is investigating the exact effects of dingoes on the fox and cat populations to determine the benefits of keeping the dog in certain areas of Australia.[78] In many areas, wild dogs live together with the most species of quolls[citation needed], except for the eastern quoll, which is probably extinct on the mainland, so wild dogs are not regarded as a threat to them.

Additionally, the disappearance of dingoes might increase the prevalence of kangaroo, rabbit and turkey[clarification needed] numbers. In the areas outside the Dingo Fence, the number of dingoes and emus is lower than in the areas inside. However, the numbers changed depending on the habitat. Since the environment is the same on both sides of the fence, the dingo was assumed to be a strong factor for the regulation of these species.[clarification needed][79] Therefore, some people demand that dingo numbers should be allowed to increase or dingoes should be reintroduced in areas with low dingo populations to lower the pressure on endangered populations of native species and to reintroduce them in certain areas. In addition, the presence of the Australian brushturkey in Queensland increased significantly after dingo baiting was conducted.[80]

Cultural impact[edit]

Cultural opinions about the dingo are often based on its perceived "cunning", and the idea that it is an intermediate between civilisation and wildness.[81]

Some of the early European settlers looked on dingoes as domestic dogs, while others thought they were more like wolves. Over the years, dingoes began to attack sheep, and their relationship to the Europeans changed very quickly: they were regarded as devious and cowardly, since they did not fight bravely in the eyes of the Europeans, and vanished into the bush.[82] Dingoes were seen as predators that killed wantonly, rather than out of hunger (similar claims are made today concerning dingo-hybrids).[83] Additionally, they were seen as promiscuous or as devils with a venomous bite or saliva, and so they could be killed unreservedly. Over the years, dingo trappers gained some prestige for their work, especially when they managed to kill hard to catch dingoes. Dingoes were associated with thieves, vagabonds, bushrangers and parliamentary opponents. From the 1960s, politicians began calling their opponents "dingo," meaning they were cowardly and treacherous, and it has become a popular form of attack since then.[56] Today, the word "dingo" still stands for "coward" and "cheat," with verb and adjective forms used, as well.[81]

The image of the dingo now ranges from the romantic to the demonic.[84][85] While some Australians see the dingo as a wild dog, others see them as slightly tame wolves, and cultural biases about each of these animals affect general perceptions about dingoes. The dingo is considered an icon of Australia, which should be preserved (at least in its "pure" form), and its possible "extinction" is compared to that of the thylacine.[3] Where dingoes are regarded as pests regardless of their "rehabilitation,"[85] this attitude can degenerate into full hatred. In the process, dingoes are sometimes considered detrimental to society and the environment (for example, that they are in general the cause for the extinction of native animals). Dingoes, whether "pure" or not, are then treated as a scourge that must be eradicated. In such cases, it is deemed acceptable to kill all wild dogs if it would save one human life.[73] In addition, there is the opinion among bureaucrats that wild dogs are cruel towards sheep and cattle and, therefore, every cruelty against them is justified.[86]

Traditionally, dogs have a privileged position in the Aboriginal cultures of Australia (which the dingo may have adopted from the thylacine), and the dingo is a well-known part of rock carvings and cave paintings.[5] Ceremonies (like a keen at the Cape York Peninsula in the form of howling[51]) and dreamtime stories are connected to the dingo, which were passed down through the generations. There are strong feelings that dingoes should not be killed[citation needed] and, in some areas, women breastfeed young cubs. In most cases, dingoes are treated with extraordinary indulgence[citation needed], although the reasons for this might not be kindness, since dogs are sometimes treated quite brutally. Nonetheless, there seems to be a big feeling of community, although the reasons seem to be clear.[clarification needed]

Similar to how Europeans acquired dingoes, the Aboriginal people of Australia acquired dogs from the immigrants very quickly. This process was so fast that Francis Barrallier (the first European to explore the Outback) discovered in 1802 that five dogs of European origin were there before him.[56] One theory holds that other domestic dogs will adopt the role of the "pure" dingo.[84] In fact, the majority of the myths about dingoes simply call them "dogs" (whether that role was adopted, or whether there was no difference for the storyteller, is unknown),[5] and other introduced animals, such as the water buffalo and the domestic cat, have been adopted into the indigenous Aboriginal culture in the forms of rituals, traditional paintings and dreamtime stories.[81]

The dingo is connected to holy places, totems, rituals and dreamtime characters. There are stories that dogs can see the supernatural, serve as guard dogs, and warn against evil powers. There is evidence that dogs have been buried with their owners to protect them against evil even after death.[87] Most of the published myths originate from the Western Desert and show a remarkable complexity. In some stories, dingoes are the central characters, while in others, they are only minor ones. One time, it is an ancestor from the dreamtime who created humans and dingoes or gave them their current shape. There are stories about creation, socially acceptable behaviour, and explanations why some things are the way they are. There are myths about shapeshifters (human to dingo or vice versa), "dingo-people," and the creation of certain landscapes or elements of those landscapes, like waterholes or mountains.

In other stories, the dingo is responsible for death. In some myths, advice and warnings are given to those who do not want to follow the social rules. Stories can show the borders of one's territory or the dingo in it might stand for certain members of the community; for example, rebellious dingoes stand for "wild" members of the tribe. The dingo has a wild and uncontrollable face in other stories, and there are many tales about dingoes that kill and eat humans (for example, the Mamu, which catches and devours the spirit of every child who roams too far from the campfire).[5] Other stories tell of a giant devil dingo, from which ordinary dingoes originate.

The dog is thereby depicted as a homicidal, malicious creature that—apart from the lack of a subtle mind—is similar to a trickster, since it plays the role of a mischievous adversary for other mythological beings. Many of them fall victim to blood-thirsty dogs or escape them. Here, individual beings have a significant meaning or sometimes become part of the landscape. The actions of these dogs result, for instance, in the creation of stones and trees from flying bones and meat or ochre from the spilled blood.[87]

Economic impact[edit]

Wild dogs are responsible for a wide range of negative and undesired impacts on the livestock industry of Australia, and they have been regarded as pests since the start of the European livestock industry. Sheep are the most frequent prey, followed by cattle and goats. Research on the real extent of the damage, though, and the reason for this problem, only started recently. Livestock can die from many causes and, when the carcass is found, it is often difficult to determine with certainty the cause of death. Since the outcome of an attack on livestock depends to a high degree on the behaviour and experience of the predator and the prey, only direct observation is certain to determine whether an attack was by dingoes or another domestic dog. Even the existence of remnants of the prey in the scat of wild dogs do not prove they are pests, since wild dogs also eat carrion. Exact numbers or reliable estimates of the damage caused by wild dogs are, therefore, hard to obtain and are seldom reliable. Even if livestock is not a big part of the dingo's diet, the extent of damage dingoes could potentially cause to the livestock industry could be much larger because of wanton killing.

The significance of dingoes as a pest is based primarily on the predation of sheep and, to a lesser extent, on cattle, and is not connected only to the direct loss of livestock. Sheep of every age are susceptible to dingo attacks, but in the case of cattle, only the calves are susceptible.[citation needed] Harassment of sheep can cause a less optimal use of grassland and miscarriages.

Distribution of wild dogs and livestock (after Breckwoldt 1988, Corbett 1995a, Fleming 1996a)

The cattle industry can tolerate low to moderate, and sometimes high, grades[clarification needed] of wild dogs (therefore dingoes are not so easily regarded as pests in these areas). In the case of sheep and goats, a zero-tolerance attitude is common. The biggest threats are dogs that live inside or near the paddock areas. The extent of sheep loss is hard to determine, due to the wide pasture lands in some parts of Australia. The numbers of cattle losses is much more variable and less well-documented. Although the loss of cattle can rise up to 30%,[50] the normal loss rate is about 0–10%.[88]

Therefore, factors such as availability of native prey, as well as the defending behaviour and health of the cattle, play an important role in the number of losses. A study in Central Australia in 2003 confirmed that dingoes only have a low impact on cattle numbers when a sufficient supply of other prey (such as kangaroos and rabbits) is available. In some parts of Australia, it is assumed that the loss of calves can be minimised if horned cattle are used instead of polled.[12] The precise economic impact is not known in this[which?] case, and it is unlikely that the rescue of some calves compensates for the necessary costs of control measures. Calves usually suffer less lethal wounds than sheep due to their size and the protection by the adult cattle, and therefore have a higher chance of surviving an attack. As a result, the evidence of a dog attack may only be discovered after the cattle have been herded back into the enclosure,[clarification needed] and signs such as bitten ears, tails and other wounds are discovered.

The opinions of cattle owners regarding dingoes are more variable than the those of sheep owners. Some cattle owners believe that it is better that the weakened mother loses her calf in times of drought so that she does not have to care for her calf, too. Therefore, these owners are more hesitant to kill dingoes.[51] The cattle industry may benefit from the predation of dingoes on rabbits, kangaroos and rats. Furthermore, the mortality rate of calves has many possible causes, and it is difficult to discriminate between them. The only reliable method to document the damage would be to document all pregnant cows, then observe their development and that of their calves.[50] The loss of calves in observed areas where dingoes were controlled was higher than in other areas. Loss of livestock is, therefore, not necessarily caused by the occurrence of dingoes and is independent from wild dogs.[89]

Domestic dogs are the only terrestrial predators in Australia that are big enough to kill fully-grown sheep, and only a few sheep manage to recover from the severe injuries. In the case of lambs, death can have many causes apart from attacks by predators, which are blamed for the deaths because they eat from the carcasses. Although attacks by red foxes are possible, such attacks are more rare than previously thought.[89] The fact that the sheep and goat industry is much more susceptible to damage caused by wild dogs than the cattle industry is mostly due to two factors: the flight behaviour of the sheep and their tendency to flock together in the face of danger, and the hunting methods of wild dogs, along with their efficient way of handling goat and sheep.

Therefore, the damage to the livestock industry does not correlate to the numbers of wild dogs in an area (except that there is no damage where no wild dogs occur[89]). Even if there are only a few wild dogs in an area, the damage to the sheep industry can be very high, since surplus killing can occur. Sometimes, extreme losses of livestock are reported (once reportedly 2,000 sheep in one night[72]) and are supposed[who?] to be increasing.

According to a report from the government of Queensland, wild dogs cost the state about $30 million annually due to livestock losses, the spread of diseases and control measures. Losses for the livestock industry alone were estimated to be as high as $18 million.[50] In Barcaldine, Queensland, up to one-fifth of all sheep are killed by dingoes annually, a situation which has been described as an "epidemic".[90] According to a survey among cattle owners in 1995, performed by the Park and Wildlife Service, owners estimated their annual losses due to wild dogs (depending on the district) to be from 1.6% to 7.1%.[91]

Despite the variety of estimates, there is little doubt that predation by dingoes can cause enormous economic damage, especially in times of drought when natural prey is sparse and the number of dingoes is still relatively high. Furthermore, wild dogs are involved in the spread of echinococcosis among cattle and sheep, as well as heartworms and parvoviruses among dogs under human care[citation needed]. An infection with echinococcosis can lead to confiscation of 90% of the intestines[clarification needed], which further leads to a value decrease of the meat and high economical damage. Furthermore, bitten livestock can only be sold for a lower price[citation needed].

Dogs are regarded as a delicacy in East Asia and Oceania, and are regularly killed for eating. In the northeast of Thailand, about 200 dingoes are killed per week to be sold on the meat market. Before the start of the 20th century, dingoes were eaten by indigenous Australians, but there are no recent reports about this practice.[6] Among the indigenous Australians, dingoes were also used as hunting aids, living hot water bottles and camp dogs. Their scalps were used as a kind of currency, their teeth were traditionally used for decorative purposes, and their fur for traditional costumes. In some parts of Australia, premiums are paid for dingo fur and scalps. The fur of dingoes generally has only a low value, and export of this fur is forbidden in states where they are protected. There is no widespread commercial catching and killing of dingoes for the purposes of obtaining their fur.

Sometimes "pure" dingoes are important for tourism, when they are used to attract visitors. However, this seems to be common only on Fraser Island, where the dingoes are extensively used as a symbol to enhance the attraction of the island. Tourists are drawn to the experience of personally interacting with dingoes. Pictures of dingoes appear on brochures, many websites and postcards advertising the island.[37] The use of dingo-urine as a repellent against dingoes and wallabies has been considered, but has not yet been economically implemented.[92]

Legal status[edit]

Until 2004, the dingo was categorized as of "least concern" on the Red List of Threatened Species. However, it has since been recategorized as "Vulnerable," following the decline in numbers to around 30% of "pure" dingoes, due to crossbreeding with domestic dogs.[1] The dingo is regarded as a regulated, but not threatened, native species under the Environment Protection and Biodiversity Conservation Act 1999 in the Commonwealth of Nations and is, therefore, protected in the national parks of the Commonwealth, as well as in World Heritage Sites and other conservation areas. However, this law also allows that dingoes can be controlled in areas where they have a proven impact on the environment. The law forbids the export of dingoes or their body parts from Australia, except for cases where it is regulated by the law. The legal status of the dingo and other wild dogs varies across the Australian federal states and territories.[50]

  • Northern Territory: The dingo is regarded as protected, not threatened, and native (due to its ecological impact) under the Territory Parks and Wildlife Conservation Act (2000). Dingoes in the Northern Territory are regarded as having an important conservational value, since interbreeding of dingoes and other domestic dogs is low in the area. However, dingoes can be legally killed when they are a danger to the livestock industry.
  • Western Australia: Dingoes and their hybrids are regarded as declared animals under the Agriculture and Related Resources Protection Act (1976). Populations must be controlled, and dingoes can be kept as pets under certain conditions. Control measures are strictly confined to livestock areas, and other domestic dogs are controlled in general. Dingoes are also regarded as unprotected native fauna under the Western Australian Wildlife Conservation Act. Although not protected, dingoes are normally not hunted without permission in conservation areas.
  • South Australia: Dingoes and their hybrids are considered pests in the sheep areas south of the Dingo Fence under the Animal and Plant Control Board (Agricultural Protection and Other Purposes) Act (1986). There, they must be controlled and can only be kept in captivity by authorised zoos and wildlife parks. North of the Dingo Fence, dingoes are regarded as legitimate wildlife and, although they are not protected, they are given a certain protection in a buffer zone of 35 km north of the Dingo Fence.
  • Queensland: Dingoes and their hybrids are regarded as pests under the Land Protection (Pest and Stock Route Management) Act 2002. All landowners are legally committed to reduce the number of all wild dogs on their lands. The dingo is regarded as wildlife and native wildlife under the Nature Conservation Act 1992 and is a natural resource (therefore protected) in conservation areas. Outside of these areas, dingoes are not regarded as native Australian and are not protected. Dingoes and their hybrids can only be kept in wildlife parks and zoos with ministerial agreement.
  • New South Wales: The Rural Lands Protection Act (1998) allocates wild dogs the status of pests, and demands from landowners that dingoes shall be decimated or eradicated.[citation needed] Although dingoes are not regarded as protected under the National Parks and Wildlife Act 1974, they are granted full protection in national parks. The dingo is regarded as a native species under the Threatened Species Conservation Act (1995), since these dogs had established populations before the European colonisation. The Wild Dog Destruction Act (1921) includes dingoes in its definition of "wild dogs". This law only affects the western part of the state, where landowners are committed to control wild dogs. The law forbids the ownership of dingoes in that region, except with legal permission. In other parts of the federal state, dingoes can be kept as pets under the Companion Animals Act (1998).
  • Australian Capital Territory: Dingoes are regarded as protected under the Nature Conservation Act (1980). On private land, the killing of wild dogs is allowed when with permission from the territory.
  • Victoria: Wild dogs are regarded as established pests under the Catchment and Land Protection Act (1994), and landowners (except from the Commonwealth) have the legal duty to hinder the spreading of wild dogs on their lands and eradicate them as much as possible. The term "wild dogs" here includes all dingoes, feral domestic dogs, dogs who became wild and crossbreeds (except for recognised breeds such as the Australian Cattle Dog).[93] The Domestic (Feral and Nuisance) Animal Act (1994) requires every dog owner to have their dogs under control at all times. The dingoes are granted a certain protection in areas that are managed by the National Parks Act (1975). Since 1998, it is possible to own dingoes as pets.[56] There is the possibility that "pure" dingoes may become officially classified as a protected species, according to official statements, and would not stand in conflict with control measures against wild dogs.[94] Update: In 2008, Dingoes were officially declared a threatened species (in danger of extinction) and are now protected.
  • Tasmania: The import of dingoes to Tasmania is forbidden under the National Parks and Wildlife Act (1970). The control of dogs that attack livestock is managed under the Dog Control Act (1987).

Control measures[edit]

See also: Dingo scalping

Dingo attacks on livestock led to widescale efforts to repel them from areas with intensive agricultural usage, and all states and territories have enacted laws for the control of dingoes.[12] In the early 20th century, fences were erected to keep dingoes away from areas frequented by sheep, and a tendency to routinely eradicate dingoes developed among some livestock owners. Established methods for the control of dingoes in sheep areas entailed the employment of specific workers on every property. The job of these people (who were nicknamed "doggers") was to reduce the number of dingoes by using steel traps, baits, firearms and other methods. The responsibility for the control of wild dogs lay solely in the hands of the landowners. At the same time, the government was forced to decimate the number of dingoes that came from unoccupied areas or reserves that might have ultimately travelled to industrial areas. As a result, a number of measures for the control of dingoes developed over time. It was also considered that dingoes travel over long distances to reach areas with richer prey populations, and the control methods were often concentrated along "paths" or "trails" and in areas that were far away from sheep areas. All dingoes were regarded as a potential danger and were hunted.

Apart from the introduction of 1080 (extensively used for 40 years and nicknamed "doggone"), the methods and strategies for decimating wild dogs have changed little over time. Caution is used at least in some control areas today, since dingoes are fully protected there. Information concerning cultural importance to indigenous people and the importance of dingoes and the impact of control measures on other species is also lacking in some areas. Historically, the attitudes and needs of indigenous people were not taken into account when dingoes were controlled. So-called dingo conservation zones are regarded as a possible solution for this problem, and these zones would mainly be based on holy dingo sites and dreamtime paths. Other factors that might be taken into account are the genetic status (degree of interbreeding) of dingoes in these areas, ownership and land usage, as well as a reduction of killing measures to areas outside of the zones. Land owners are increasingly committed to regularly record where individual dingoes and their tracks are most frequent and cause the most damage. Also, birth, damage and mortality rates of livestock should be recorded. However, most control measures and the appropriate studies are there to minimise the loss of livestock and not to protect dingoes. In areas of cattle industries, there are few or no control measures,[citation needed] and efforts are mostly limited to occasional shootings and poisonings. Government controlled use of 1080 is performed only every third year, when field observations prove the claims of high livestock losses and dingo numbers.

Increasing pressure from environmentalists against the random killing of dingoes, as well as the impact on other animals, demanded that more information needed to be gathered to prove the necessity of control measures and to disprove the claim of unnecessary killings.Today, permanent population control is regarded as necessary to reduce the impact of all wild dogs and to ensure the survival of the "pure" dingo in the wild.[50]

A part of the dingo fence

Trapping[edit]

Trapping and removal is an essential part of the control measures in the highlands of southeastern New South Wales and Northern Victoria. Where steel traps and baits cannot, or are not, allowed to be used (for example, residential zones), cage traps are used.

Deterrence[edit]

One method that does not have any proven effect is to hang dead dogs along the borders of the property in the belief that this would repel wild dogs.[12]

Guardian animals[edit]

To protect livestock, livestock guardian dogs (for example, Maremmas), donkeys, alpacas and llamas are used.[95][96]

To keep wild dogs away from certain areas, efforts are taken to make these areas unattractive for them (for example, by getting rid of food waste) and therefore forcing them to move elsewhere. Control through deliberately spreading disease is normally not considered. Such attempts probably would not be successful, because typical dog diseases are already present in the population. Additionally, dogs under human care would also be susceptible. Other biological control methods are not regarded as achievable, since there would be a high risk of decimating dogs under human care.

Shooting[edit]

It does happen that dingoes are hunted and shot by people on horseback, or that a premium is sold for shot dingoes.

Dingo Fence[edit]

In the 1920s, the Dingo Fence was erected on the basis of the Wild Dog Act (1921) and, until 1931, thousands of miles of Dingo Fences had been erected in several areas of South Australia. In the year 1946, these efforts were directed to a single goal, and the Dingo Fence was finally completed. The fence connected with other fences in New South Wales and Queensland. The main responsibilities in maintaining the Dingo Fence still lies with the landowners, whose properties border on the fence and receive financial support from the government.

Reward system[edit]

A reward system (local, as well from the government) was active from 1846 to the end of the 20th century, but there is no evidence that – despite the billions of dollars spent – it was ever an efficient control method. Therefore, its importance declined over time.[6]

Poisoning[edit]

Warning of poisonous sodium fluoroacetate baits
Dead wild dogs that have been hung on a fence

Strychnine is still used in all parts of Australia.

Baits with the poison 1080 are regarded as the fastest as safest method for dog control, since they are extremely susceptible. Even small amounts of poison per dog are sufficient (0.3 mg per kg).[50] The application of aerial baiting is regulated in the Commonwealth by the Civil Aviation Regulations (1988). The assumption that the Tiger Quoll might be damaged by the poison led to the dwindling of areas where aerial baiting could be performed. In areas where aerial baiting is no longer possible, it is necessary to put down baits.

Over the last years, cyanide-ejectors and protection collars (filled with 1080 on certain spots) have been tested.[97][98]

The eradication of dingoes due to livestock damage decreased along with the importance of the sheep industry and the usage of strychnine (which beforehand had been used for 100 years) in the 1970s. The number of doggers also decreased and the frequency of government-approved aerial baiting increased. During this period, many farmers in Western Australia switched to the cattle industry, and findings in the area of biology led to a significant change in control measures and techniques in association with reduced costs and increased efficiency. At the same time, the importance of 1080 increased, and the first anxieties arose that the number of dingoes might have decreased so much that they may become locally extinct. Observations on the ecology of dingoes led to the practice of placing baits near water holes, hiding places and prey sites.

Spaying and neutering[edit]

Owners of dingoes and other domestic dogs are sometimes asked to spay or neuter their pets and keep them under observation to reduce the number of stray/feral dogs and prevent interbreeding with dingoes (for instance under the Territory Parks and Wildlife Conservation Act (2000)).[50]

Efficiency of measures[edit]

The efficiency of control measures was questioned in the past and is often questioned today, as well as whether they stand in a good cost-benefit ratio. The premium system proved to be susceptible to deception and to be useless on a large scale, and can therefore only be used for getting rid of "problem-dogs".[12][99] Animal traps are considered inhumane and inefficient on a large scale, due to the limited efficacy of baits. Based on studies, it is assumed that only young dogs that would have died anyway can be captured.[52] Furthermore, wild dogs are capable of learning and sometimes are able to detect and avoid traps quite efficiently. In one case, a dingo bitch followed a dogger and triggered his traps one after another by carefully pushing her paw through the sand that covered the trap.[82]

Poisonous baits can be very effective when they are of good meat quality; however, they do not last long[100] and are occasionally taken by red foxes, quolls, ants and birds. Aerial baiting can nearly eliminate whole dingo populations.[52] Livestock guardian dogs can effectively minimise livestock losses, but are less effective on wide open areas with widely distributed livestock. Furthermore, they can be a danger to the livestock or be killed by control measures themselves when they are not sufficiently supervised by their owners.[98] Fences are reliable in keeping wild dogs from entering certain areas, but they are expensive to build, need permanent maintenance, and only cause the problem to be relocated.

According to studies, control measures can eliminate 66% to 84% of a wild dog population, but the population can reach its old numbers very quickly over the course of a year, depending on the season, such as by immigration of young dogs from other areas. Only a cohesive coordinated control in all areas could be efficient in the long run, if at all.[71] Control measures mostly result in smaller packs respectively in a disruption of the pack structure. The measures seem to be rather detrimental to the livestock industry because the empty territories are taken over by young dogs and the predation then increases. Nonetheless, it is regarded as unlikely that the control measures could completely eradicate the dingo in Central Australia, and the elimination of all wild dogs is not considered a realistic option.

Conservation[edit]

Dingoes are reasonably abundant in large parts of Australia, but there is some argument that they are endangered due to interbreeding with other dogs in many parts of their range.[1] Dingoes are not a protected species, but they are regulated under federal law and, thus, their status varies in different states and territories. Dingoes receive varying levels of protection in conservation areas such as national parks and natural reserves in New South Wales, the Northern Territory and Victoria, Arnhem Land and other Aboriginal lands, UNESCO World Heritage Sites, and the whole of the Australian Capital Territory. In some states, dingoes are regarded as declared pests and landowners are allowed to control the local populations. Throughout Australia, all other wild dogs are considered pests.

Dingo with a tagged ear on Fraser Island

The dingoes of Fraser Island are considered to be of significant conservational value. Due to their geographic and genetic isolation, they are considered to be the most similar to the original dingoes, and they are seen as the most pure dingo population. The dingoes there are not "threatened" by interbreeding with other domestic dogs. Because of their conservational value, outrage was sparked in January 2013 when two six-month-old dingo pups were found dead, believed to have been run over near Lake McKenzie.[101] The couple who found the dingoes were outraged at the reaction of the rangers, and Fraser Coast area manager Ross Belcher said that there will be serious penalties for those who kill or injure Fraser Island dingoes.[102]

In February 2013, a report on Fraser Island dingo management strategies was released, with options including ending the intimidation of dingoes, tagging practice changes and regular veterinarian checkups, as well as a permanent dingo sanctuary on the island.[103]

Groups that have devoted themselves to the conservation of the "pure" dingo by using breeding programs include the Australian Native Dog Conservation Society and the Australian Dingo Conservation Association. Presently, the efforts of the dingo conservation groups are considered to be ineffective because most of their dogs are untested or are known to be hybrids.[6]

Dingo conservation efforts focus primarily on preventing interbreeding between dingoes and other domestic dogs in order to conserve the population of pure dingoes. This is extremely difficult and costly. Conservation efforts are hampered by the fact that it is not known how many pure dingoes still exist in Australia. Steps to conserve the pure dingo can only be effective when the identification of dingoes and other domestic dogs is absolutely reliable, especially in the case of living specimens. Additionally, conservation efforts are in conflict with control measures.

Conservation of pure and survivable dingo populations is promising in remote areas, where contact with humans and other domestic dogs is rare. Under New South Wales state policy in parks, reserves and other areas not used by agriculture, these populations are only to be controlled when they pose a threat to the survival of other native species. The introduction of "dog-free" buffer zones around areas with pure dingoes is regarded as a realistic method to stop interbreeding. This is enforced in the way that all wild dogs can be killed outside of the conservation areas. However, studies from the year 2007 indicate that even an intensive control of core areas is probably not able to stop the process of interbreeding.[104]

There is presently no information regarding what opinions the public has regarding the conservation of dingoes. There is no unity on the definition of "pure" dingoes and to what extent they should be controlled.[105]

As a pet and working dog[edit]

Opinion is divided about the keeping of dingoes as pets or as working dogs. Some consider the dingo unsuitable for domestication, while others see a domesticated dingo as no different from any other domesticated dog.[106] In this vein, dingoes would have the right to be recognised as a dog breed, and domestication would be the only reliable way of ensuring the survival of the "pure" dingo.[107] Some disagree that the dingo should be labeled a dog breed, as they believe "true" dingoes can be tamed but not truly domesticated.[108]

Dingoes are sometimes kept as pets, although their tendencies as wild animals are difficult to suppress.

Dingoes can be very tame when they come in frequent contact with humans.[24] Furthermore, some dingoes live with humans (due to practical, as well as emotional reasons). Many indigenous Australians and early European settlers lived alongside dingoes. Indigenous Australians would take dingo pups from the den and tame them until sexual maturity and the dogs would leave.[108] Alfred Brehm reported cases where dingoes that were completely tame and, in some cases, behaved exactly like other domestic dogs (one was used for shepherding heavy livestock), as well as specimens that remained wild and shy. He also reported about dingoes that were aggressive and completely uncontrollable, but he was of the opinion that these reports "should not get more attention than they deserve," since the behaviour depends on how the dingo was raised since early puppyhood. He believed that these dogs could become very decent pets.[26]

According to Eberhard Trumler,[who?] dingoes are very smart and affectionate. To would-be owners, he recommended the provision of a large escape-proof enclosure and a partner of the opposite sex. During heat, dingoes are harder to manage than other domestic dogs which, combined with their attachment to their owners, can lead to problems, since they want to follow their owners and never miss the opportunity to feed. Dingoes are supposed[who?] to find every weak spot of an enclosure or residence, escape for a while and stray through towns and villages. Their intellectual ability is supposedly[who?] connected to an enormous ability to learn and a lightning perception. Dingoes have a reputation for not handling pressure, but this conflicts with their record as working dogs. They are suitable as shepherd dogs, appearing to see a purpose in it (keeping together a familiar group is in their nature) and, even today, some dingoes are employed as shepherd dogs. In addition, dingoes have strong toileting instincts and can be easily housebroken.

In 1976, the Australian Native Dog Training Society of NSW Ltd. was founded. Until this time, the ownership of dingoes was illegal.[citation needed] In mid-1994, the Australian National Kennel Council(ANKC) officially recognised the dingo as Australia's national dog breed, and a breed standard was published some years later. The dingo is listed in Group 4 (hound) of the ANKC.[109] However, it is still illegal in some states to own, breed or sell dingoes, as it is in some countries.[15]

In South Australia, dingoes can only be kept in specially licensed zoos, circuses and authorized research institutions. South Australia is a particularly sensitive region, due to extensive sheep farming conflicting with large populations of wild dingoes in the north of the state. Dingoes are bred by certain clubs and private individuals in Australia and the United States. The dingo is not regarded as a dog breed by the Fédération Cynologique Internationale. However, the American Rare Breed Association (ARBA) regards the dingo as a breed belonging to the Spitz and Primitive Group.

Goals[edit]

In 1998 in New South Wales, the dingo was reclassified as a pet in order to save the species from extinction. Breeding programs were introduced, which were considered to be the best option available for safeguarding the continuation of the species, with the goal of returning them to the wild at a later date.[106]

Dingoes have also been bred for sale or use as working dogs. The use of dingoes at customs was first attempted in 1976 in Victoria. However, some people speculated that these dogs were cross-breeds of dingoes and shepherd dogs.[15]

Criticism[edit]

The ownership of dingoes as pets and their breeding is widely criticized. The main criticism is that the activities and the resulting consequences of the dingo conservation groups, "dingo farms" and legislation for legal ownership of dingoes for people in public, is seen to be an additional threat to the survival of the pure dingoes. This fear exists because the majority of these breeding activities effectively expedite the interbreeding of dingoes and other domestic dogs, when the identification of a pure dingo is not absolutely correct respectively when hybrids are sold as "pure" dingoes.[6][clarification needed]

Supporters of breeding programmes are only mildly optimistic about a successful outcome. Success in the form of a population viable for future re-wilding cannot be easily accomplished.[110] According to David Jenkins,[who?] the breeding and reintroduction of pure dingoes is no easy option and, at the time[when?], there were no studies that seriously dealt with this topic, especially in areas where dingo populations are already present.[111]

An additional threat is that breeders may unconsciously select tamer dingoes by breeding individuals who are easier to manage. Therefore it may happen that, over the years, the tame populations may become less suitable for living in the wild than their ancestors. In addition, a loss of genetic diversity (thus resulting in a higher susceptibility to diseases) might occur due to a small founding population, and negative changes could occur simply because the dogs were captive-bred. Furthermore, some features that are necessary for survival in the wild might "fade" under the conditions of domestication (for example, hunting techniques) because they are no longer needed.

Another criticism is that adult dingoes are viewed by some[who?] to be unsuitable as pets in the same ways as other domestic dogs. Dingoes are regarded[who?] as more independent-minded than other domestic dogs, making domestication reportedly[who?] difficult. As dingoes age, they succumb[clarification needed] to their natural instincts and become more likely to escape into the wild.[106] Furthermore, most people are unable to provide a dingo with what it needs, and dingoes may not react positively to domestication and training. Supposedly, only few dingoes and dingo-hybrids would reach an old age, since the owners would not know how to handle them. When a dingo is not socialised, it would be hard to control and develop behavioural problems from aspects of domestic life more easily tolerated by other dog breeds. To make dingoes more suitable as lapdogs, breeders would need to cross them with other domestic dogs.[112]

Interbreeding with domestic dogs[edit]

Although dingo-like, this wild dog has an atypical colouration and is therefore most likely a dingo-crossbreed.

European domestic dogs first arrived in Australia during the European colonisation. These dogs reverted to the wild (both unintentionally and intentionally), produced feral populations and interbred with the existing dingoes. Hybrids of dingoes and domestic dogs exist today in all wild dog populations of Australia, with their numbers having increased to such a degree that any completely "pure" populations may no longer exist.[56] The degree of interbreeding is locally so high, for instance in urban and rural areas, that there are big populations consisting purely of hybrids. Estimates from the 1990s assumed a proportion of dingo-hybrids of about 78% in the wild.[113] It is not clear how large the current population of hybrids is today.

Dingo-like domestic dogs and dingo-hybrids can be generally distinguished from "pure" dingoes by their fur colour, since there is a wider range of colours and patterns among them than among dingoes. In addition, the more dog-typical kind of barking exists among the hybrids, and differences in the breeding cycle,[114] certain skull characteristics,[115] and genetic analyses[116] can be used for differentiation. Despite all the characteristics that can be used for distinguishing between dingoes and other domestic dogs, there are two problems that should not be underestimated. First, there is no real clarity regarding at what point a dog is regarded as a "pure" dingo,[105] and, secondly, no distinguishing feature is completely reliable—it is not known which characteristics permanently remain under the conditions of natural selection.

In science, there are two main opinions regarding this process of interbreeding. The first, and likely most common, position states that the "pure" dingo should be preserved via strong controls of the wild dog populations, and only "pure" or nearly "pure" dingoes should be protected.[117] The second position is relatively new and is of the opinion that people must accept that the dingo has changed and that it is impossible to bring the "pure" dingo back. Conservation of these dogs should therefore be based on where and how they live, as well as their cultural and ecological role, instead of concentrating on precise definitions or concerns about "genetic purity".[118] Both positions are controversially discussed.

There is a wider range of fur colours, skull shapes and body size in the modern-day wild dog population than in the time before the arrival of the Europeans. Over the course of the last 40 years ,[when?] there has been an increase of about 20% in the average wild dog body size.[119] It is currently unknown whether, in the case of the disappearance of "pure" dingoes, remaining hybrids would alter the predation pressure on other animals. It is also unclear what kind of role these hybrids would play in the Australian ecosystems. However, it likely that the dynamics of the various ecosystems will not be disturbed by this process.[12]

Attacks on humans[edit]

Main article: Dingo attack

Although dingoes are large enough to be dangerous, they generally avoid conflict with humans. Apart from the well-known case in which an infant was taken from a campsite (see below), there have been numerous confirmed dingo attacks, often involving people feeding wild dingoes, particularly on Fraser Island, a special center of dingo-related tourism (see main article). Most dingo attacks are minor in nature, but some can be major, and a few can be fatal. Many Australian national parks have signs advising visitors not to feed wildlife, partly because this practice is not healthy for the animals, and partly because it may encourage undesirable behaviour, such as snatching or biting by dingoes, goannas and some birds.

Azaria Chamberlain dingo attack[edit]

On 17 August 1980, a nine-week-old girl named Azaria Chamberlain was taken by a dingo near Uluru (Ayers Rock) and killed.[32] Her mother, Lindy Chamberlain, was suspected and wrongly convicted of murder, as the court did not believe that an animal generally shy of humans would be capable of such an act. After serving more than three years of her sentence, Lindy was released from prison when the jacket of the baby was found in a dingo den. The mother was thereafter found innocent, but the cause of death was not officially listed as a dingo attack until 12 June 2012.[120][121][122][123]

Fraser Island attacks[edit]

  • In April 1998, a 3-year-old Norwegian girl was bitten and scratched by a dingo.[124]
  • On 30 April 2001, nine-year-old Clinton Gage was attacked and killed by two dingoes near Waddy Point on Fraser Island. The incident and the resultant culling of 31 dingoes caused a large outcry among the residents. There were many protests and the suggestion was made to erect fences.[125][126]
  • On 26 April 2011, a three-year-old girl was attacked on Fraser Island by two dingoes. She suffered serious puncture wounds to her leg.[127]
  • In November 2012, a six-month-old dingo by the name[who?] of Inky was killed by rangers on Fraser Island after continued aggressive and dangerous behaviour towards people. The dangerous behaviour included included "lunging" at a family, coming out of the bushland at high speed towards volleyball players, and grabbing two tourists on separate occasions with his mouth, not breaking the skin on either occasion.[128] Rangers attempted to trap the dangerous dingo for a month before they were successful. The captured animal was then euthanised.[129] One Dingo advocacy group argued that, as a juvenile, the dingo's aggressive behaviours would be considered normal for his young age.[128] Soon after, Inky's brother Byron was killed by rangers, although his documented incidents never reached the serious Code E level that his brother's had.[130]

Dingo conservation groups on Fraser Island have become frustrated with the killing of dingoes that attack humans.[citation needed] It has been proposed[who?] that problem dingoes be relocated to a wildlife sanctuary.[131] Queensland Environment Minister Andrew Powell said that the Fraser Island government should work to better educate people about dingoes to help stop attacks.[132]

Conclusions[edit]

Articles published about dingo attacks blame them on habituation, especially through the intentional and unintentional feeding of dingoes.[citation needed] The more frequently these animals are fed or allowed to scavenge on waste food, the more likely they are to react aggressively towards humans when they no longer receive or find food. It is further thought[who?] that dingoes might have started to regard the food sources found (garbage cans, leftovers and handouts) as part of their territory. Attacks then occur with humans seen as competition, and dingoes simply reacting to protect their food supply.

Even when habituation to humans seems to be the general cause for attacks, it is not absolutely clear, and therefore the overall threat towards people is not known for sure. Some attacks might result from the "play" of young cubs, especially with children. Attacks can also be caused by mistaken reactions of humans to aggressive and dominant behaviour of dingoes. That some dingoes might regard humans as prey is a possibility, as children or incapacitated adults could be theoretically overpowered.[37][126] Dr. Bradley Smith[who?] said that Fraser Island has a problem with humans and not with the dingoes, that dogs who were labelled "aggressive" were simply behaving naturally.[133]

The behaviour of humans might undermine efforts to guard against dingo attacks. Therefore, the change in human behaviour is at the centre of attention.[who?] Warning signs like "Beware of Dingoes" seem to have lost their effect on Fraser Island, despite the high number of such signs.[citation needed] Furthermore, some humans[which?] do not realise how adaptive and quick dingoes are. Therefore, humans do not remain attentive enough.[citation needed] They[who?] do not consider, for instance, that dingoes steal food like fruits and vegetables. In addition, some tourists seemed to be confused by the high number of rules in some parks, and they have been prompted in some cases to actively feed the wild animals.[37][50][125][126]

Problems in classification[edit]

Borneo-dogs, which look very similar to Australian dingoes

There is no general agreement (scientific or otherwise) regarding what the dingo is, in a biological sense, since it has been called "wolf," "dingo," "dog," and "wild dog".[85] Even within the scientific community, the dingo is given several names. There is no consensus regarding whether the dingo is a feral or native animal, or what kinds of dogs should be classed as "dingoes". Thus, some people consider the New Guinea Singing Dog, the Basenji, the Carolina Dog[56] and certain other dog populations to be dingoes. Evidence indicates a discord concerning the status of these dogs, as well.

Dingoes have been variously considered to be wild dogs,[32] the progenitor of domestic dogs,[56] the ancestor of modern dog breeds, a separate species,[134] a link between wolf and domestic dog,[56] a primitive canine species[135] or primitive domestic dog,[6] a "dog-like" relative of wolves[136] or a subspecies of the domestic dog.[137] Others consider them to be native dogs of Asia,[138] a relatively unchanged form of early domestic dog,[12] part wolf and part dog,[139] or to have been selectively bred from wolves.[95] Then again, some do not consider dingoes feral any more but completely wild, since they have been living under natural selection for a very long time.[140] According to present scientific consensus and knowledge, dingoes are domestic dogs that arrived at their present distribution with humans, adapted to the respective conditions and are no more "primitive" or "primordial" than other domestic dogs.[141]

Carolina dogs; some of these dogs bear a great similarity to dingoes.

The Australian dingo has never been subject to the artificial selection that produced modern dog breeds,[84] and it may be an undomesticated descendant of an extinct Asian wolf.[142] However, compared to the European grey wolf, dingoes have an approximately 30% lower relative brain size,[143] reduced facial expressions,[144] reduced impressive behaviour,[24] curled tails that can be carried over the back, and generally a permanent fertility in males—features that all known domestic dogs share and are considered to be caused by domestication.[19][24] It might happen that one and the same source names the dingo as a subspecies of the grey wolf, but lists all other domestic dogs as separate species.[145] Likewise, the scientific name of the dingo might be Canis lupus dingo, but the dingo is regarded as a separate species, nonetheless.[146] Alfred Brehm originally considered the dingo to be a separate species but, after examining several different specimens, he came to the conclusion that they could only be domestic dogs.[26] In contrast, William Jardine considered the dingo to be an entirely separate species, while contemporary French naturalists regarded them as feral dogs.[147] Even among modern-day scientists, dingoes and other domestic dogs are sometimes considered two separate species, despite small genetic, morphological and behavioural differences.

The phenomenon of interbreeding between both is then attributed to the statement that all wolf-like species can interbreed and produce fertile offspring.[21] However, breeding experiments in Germany could only prove an unrestricted fertility in the offspring of domestic dogs and grey wolves. Hybrids between domestic dogs and coyotes, and domestic dogs and Golden Jackals, had communication problems[clarification needed] among each other, as well to the parent species. From the third hybrid generation on, a decrease in fertility and an increase in genetic damage[clarification needed] was observed among the coyote-hybrids and jackal-hybrids.[141] Observations of this kind have never been made for hybrids of dingoes and other domestic dogs, only that dingoes and other domestic dogs can freely interbreed with each other.[148]

The choice of classification can have a direct impact on the dingo. Dingoes officially cease to exist outside of national parks and become unprotected wild dogs.[85] The term "wild dog," itself, sometimes only includes dingoes and their hybrids[149] or respectively excludes dingoes.[150] Another view is that dingoes are "only" feral outside of national parks, with this term having a more negative meaning than the term "wild".

On the other hand, dingoes have been "rehabilitated" in some way, by changing their status from pests to "Australia's native dog" or, more subtly, from a subspecies of the domestic dog to that of the grey wolf. The undertone in the Australian press seemed to be that being a grey wolf or an Asian wolf means that the dingo is more "wild" and, therefore, more desirable than a companion animal (domestic dog). It is possible that the habit of calling the dingo only "dog" (not "wild dog") in colloquial language indicates a form of familiarity or debasing. In the last case, it might be morally easier to kill a dog when it causes problems because it would not have the "high status" of a wolf or dingo.[85] Sometimes, it is considered bad that dingoes are domestic dogs, that they are descended from them and not "directly" from the grey wolf.[151] In short, if the dingo is regarded as native, then it is worthy of protection. But if it is considered to be "just" a variant of the domestic dog, it is regarded as a pest and should be eradicated.[29][72]

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  100. ^ Twigg, Laurie E.; Eldridge, Steve R.; Edwards, Glenn P.; Shakeshaft, Bernie J.; Depreu, Nicki D.; Adams, Neville (2000). "The longevity and efficacy of 1080 meat baits used for dingo control in central Australia". Wildlife Research 27 (5): 473–81. doi:10.1071/WR99044. 
  101. ^ "Pup deaths anger dingo group". abc.net.au. 9 January 2013. Retrieved 17 January 2013. 
  102. ^ Walker, Carrie (8 January 2013). "Dead dingo pups in car park distress Fraser Island tourists". Fraser Coast Chronicle. Retrieved 17 January 2013. 
  103. ^ "Dingo sanctuary considered for Fraser Island". The New Zealand Herald. 27 February 2013. Retrieved 27 February 2013. 
  104. ^ "Predation and Hybridisation by Feral Dogs (Canis lupus familiaris) – proposed key threatening process listing". New South Wales Government. 29 August 2008. Retrieved 13 May 2009. 
  105. ^ a b Brad Purcell, Robert Mulley, Robert Close (2008). "Genetic characterisation of dingoes in the Blue Mountains world heritage area". 14th Australasian Vertebrate Pest Conference. Darwin: Invasive Animals CRC. p. 140. Archived from the original on 24 April 2009. Retrieved 13 May 2009. 
  106. ^ a b c "Australia's dingo dogs face extinction". USA Today. 7 October 2003. Retrieved 14 May 2009. 
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  108. ^ a b Coppinger, Raymond and Lorna (2001). Dogs: A Startling New Understanding of Canine Origin, Behavior, & Evolution. New York: Scribner. pp. 45, 67. ISBN 0-684-85530-5. 
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  116. ^ Wilton, Alan. "Genetic Diversity in the Dingo". dingosanctuary. Archived from the original on 2004-02-19. Retrieved 14 May 2009. 
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  143. ^ Hemmer, Helmut (1983). Domestikation, Verarmung der Merkwelt (Domestication: the decline of environmental appreciation) (in german). Braunschweig: Friedr. Vieweg & Sohn Verlagsgesellschaft mbH. ISBN 3-528-08504-5. 
  144. ^ Trumler, Eberhard (1982). Ein Hund wird geboren; Der Ratgeber für den Hundefreund (in german). Muenich: R. Piper GmbH & Co. KG. ISBN 3-492-02775-X. 
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Further reading[edit]

  • J.S. Bacon: The Australian Dingo: The King of the Bush. McCarron Bird, Melbourne 1955.
  • R. Breckwoldt: A Very Elegant Animal: the Dingo. Angus and Robertson, Australia 1988.
  • Deborah Bird Rose: Dingo makes us Human, Life and Land in an Aboriginal Australian culture. Cambridge University Press, New York, Oakleigh 1992, ISBN 0-521-39269-1.
  • Chris R. Dickman: A Symposium on the Dingo. Royal Zoological Society of New South Wales, Sydney 1999, ISBN 0-9586085-2-0.
  • Erich Kolig: Aboriginal dogmatics: canines in theory, myth and dogma. In: Bijdragen tot de Taal-, Land- en Volkenkunde 134. Nr. 1, Leiden 1978, Seite 84–115.
  • Peter Fleming, Laurie Corbett, Robert Harden, Peter Thomson: Managing the Impacts of Dingoes and Other Wild Dogs. Bureau of Rural Sciences, Commonwealth of Australia, 2001.
  • Western Australian Wild Dog Management Strategy 2005. August 2005.
  • Georgette Leah Burns, Peter Howard: When wildlife tourism goes wrong: a case study of stakeholder and management issues regarding dingoes on Fraser Island. Faculty of Environmental Sciences, Griffith University.
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Gray wolf

"Wolf" redirects here. For other uses of "wolf" and "wolves", see Wolf (disambiguation).
"Gray Wolves" and "She-wolf" redirect here. For other uses, see Gray Wolves (disambiguation) and She-wolf (disambiguation).

The gray wolf[a] (Canis lupus[b]), also known as the timber wolf,[5][6] true wolf[7] or western wolf[c] is a canid native to the wilderness and remote areas of North America, Eurasia, and North Africa. It is the largest extant member of its family, with males averaging 43–45 kg (95–99 lb), and females 36–38.5 kg (79–85 lb).[9] Like the red wolf, it is distinguished from other Canis species by its larger size and less pointed features, particularly on the ears and muzzle.[10] Its winter fur is long and bushy, and predominantly a mottled gray in color, although nearly pure white, red, or brown to black also occur.[6]

The gray wolf is the most specialised member of the genus Canis, as demonstrated by its morphological adaptations to hunting large prey, its more gregarious nature,[11] and its highly advanced expressive behavior.[12][13] It is nonetheless closely related enough to smaller Canis species, such as the eastern wolf,[8] coyote[14] and golden jackal[15][16] to produce fertile hybrids. It is the only species of Canis to have a range encompassing both the Old and New Worlds,[8] and originated in Eurasia during the Pleistocene, colonizing North America on at least three separate occasions during the Rancholabrean.[8] It is a social animal, travelling in nuclear families consisting of a mated pair, accompanied by the pair's adult offspring.[17] The gray wolf is typically an apex predator throughout its range, with only humans and tigers[11][18][19][20] posing a serious threat to it. It feeds primarily on large ungulates, though it also eats smaller animals, livestock, carrion, and garbage.[6]

The gray wolf is one of the world's best known and well researched animals, with probably more books written about it than any other wildlife species.[21] It has a long history of association with humans, having been despised and hunted in most pastoral communities due to its attacks on livestock, while conversely being respected in some agrarian and hunter-gatherer societies.[22] It is the sole ancestor of the dog, which genetic and paleontological records indicate was first domesticated in Europe 18,800-32,100 years ago by hunter-gatherers.[23] Although the fear of wolves is pervasive in many human societies, the majority of recorded attacks on people have been attributed to animals suffering from rabies. Non-rabid wolves have attacked and killed people, mainly children, but this is unusual, as wolves are relatively few, live away from people, and have been taught to fear humans by hunters and shepherds.[24]

The gray wolf was once one of the world's most widely distributed mammals, living throughout the northern hemisphere north of 15°N latitude in North America and 12°N in India. However, deliberate human persecution has reduced the species' range to about one third, due to livestock predation and fear over attacks on humans. The species is now extinct in much of Western Europe, in Mexico and much of the USA. In modern times, the gray wolf occurs mostly in wilderness and remote areas, particularly in Canada, Alaska and northern USA, Europe, and Asia from about 75°N to 12°N. Wolf population declines have been arrested since the 1970s, and have fostered recolonization and reintroduction in parts of its former range, due to legal protection, changes in land-use and rural human population shifts to cities. Competition with humans for livestock and game species, concerns over the danger posed by wolves to people, and habitat fragmentation pose a continued threat to the species. Despite these threats, because of the gray wolf's relatively widespread range and stable population, it is classified as Least Concern by the IUCN.[1]

Evolution and taxonomy[edit]

Phylogenetic tree of interfertile Canis

Origins[edit]

Ancestry[edit]

The species' most likely ancestral candidate is Canis lepophagus, a small, narrow skulled North American canid of the Miocene era, which may have also given rise to C. latrans.[25] After the extinction of the large bodied Borophaginae family, C. lepophagus developed into a larger, broader-skulled animal. Fossils of this larger form of C. lepophagus found in northern Texas may represent the ancestral stock from which C. lupus derives.[26] The first true wolves began to appear at the end of the Blancan North American Stage and the onset of the early Irvingtonian. Among them was C. priscolatrans, a small species closely resembling modern-day C. rufus, which colonized Eurasia by crossing the Bering land bridge. The new Eurasian C. priscolatrans population gradually evolved into C. mosbachensis, which subsequently developed in the direction of C. lupus.[25] The earliest identifiable C. lupus remains date back to the Middle Pleistocene, and occur in Beringia.[27]

Subspeciation[edit]

MtDNA studies have shown that there are at least four extant C. lupus lineages; the most ancient is that of C. l. lupaster (native to North, West, and East Africa), which is thought to have originated as early as the Middle to Late Pleistocene.[15] All other lineages occur together in the Indian Subcontinent, the oldest of which is the Himalayan wolf (native to the Himalayan region of eastern Kashmir, Himachal Pradesh, parts of Tibet and eastern Nepal), which is thought to have originated 800,000 years ago, when the Himalayan region was going through major geologic and climatic upheaval. C. l. pallipes, likely diverged from the Himalayan wolf 400,000 years ago.[28][29] The youngest C. lupus lineage in India is represented by C. l. chanco (native to the northwestern Himalayan region of Kashmir), which originated 150,000 years ago. This last lineage, known as the Holarctic clade, expanded into Europe and North America, as shown by it sharing genetic markers with domestic dogs, European and North American wolves.[28][29]

The now extinct Japanese wolves were descended from large Siberian wolves which colonised the Korean Peninsula and Japan, before it separated from mainland Asia, 20,000 years ago during the Pleistocene. During the Holocene, the Tsugaru Strait widened and isolated Honshu from Hokkaidō, thus causing climatic changes leading to the extinction of most large bodied ungulates inhabiting the archipelago. Japanese wolves likely underwent a process of island dwarfism 7,000–13,000 years ago in response to these climatological and ecological pressures. C. l. hattai (formerly native to Hokkaidō) was significantly larger than its southern cousin C. l. hodophilax, as it inhabited higher elevations and had access to larger prey, as well as a continuing genetic interaction with dispersing wolves from Siberia.[30]

C. lupus colonized North America during the late Rancholabrean era through the Bering land bridge in at least three separate invasions, with each wave being represented by one or more different Eurasian gray wolf clades.[8] Among the first to enter was a broad-skulled, hypercarnivorous ecomorph which never expanded its range below the Wisconsin ice sheet, likely due to competitive exclusion by C. dirus populations in the south, with both dying out during the Quaternary extinction event without leaving any modern descendants.[31] The first gray wolves to permanently enter North America were the ancestors of C. l. baileyi, though these were followed and displaced by C. l. nubilus and pushed southwards. C. l. nubilus was in turn displaced from its northern range by C. l. occidentalis, likely during the Holocene, a process which may have continued into historic times.[8]

Subspecies[edit]

Globally, gray wolf taxonomy has been subject to numerous revisions, particularly in North America. As of 2005,[32] 37 subspecies of C. lupus are recognised by MSW3. Its list includes the domestic dog, dingo, eastern wolf and red wolf, but lists C. l. italicus and C. l. communis as synonyms of C. l. lupus, and C. l. lupaster as a subspecies of golden jackal.[33]

Old world gray wolf subspecies[edit]

In 1995, mammologist Robert Nowak recognized five subspecies from Eurasia based on skull morphology; C. l. lupus, C. l. albus, C. l. pallipes, C. l. cubanensis and C. l. communis.[34] In 2003, Nowak also recognized the distinctiveness of C. l. arabs, C. l. hattai, C. l. hodophilax and C. l. lupaster.[35] Furthermore, genetic studies on gray wolves in Italy revealed that, unlike several European gray wolf populations, Italian wolves do not share haplotypes with either other gray wolves or domestic dogs, and are morphologically distinct enough to be classed as a separate subspecies; C. l. italicus.[36][37][38]

Kopaliani et al. (2011) made phylogenetic comparisons of the mtDNA sequences of both gray wolves and golden jackals, and discovered that the African wolf, which was once thought to be a golden jackal, is in fact a subspecies of gray wolf, C. l. lupaster.[39]

Rueness et al. (2014) showed that wolves in the Caucasus Mountains of the putative Caucasian subspecies, C. l. cubanensis, are not genetically distinct enough to be considered a subspecies, but may represent a local ecomorph of C. l. lupus.[40]

North American gray wolf subspecies[edit]

In 1944, American zoologist Edward Goldman recognized as many as 23 subspecies in North America, based on morphology alone.[41] In 1995, mammologist Ronald Nowak disputed these classifications, based on his comparison of numerous wolf skulls from throughout the continent. He concluded that there are only five North American subspecies: C. l. occidentalis, C. l. nubilus, C. l. arctos, C. l. baileyi and C. l. lycaon. Wilson et al. (2000), a genetic study of canids from Algonquin Provincial Park, indicated that C. l. lycaon was a separate species from C. lupus, more closely related to C. rufus.[42]

In a monograph prepared within the United States Fish and Wildlife Service (USF&WS), Chambers et al. (2012) reviewed many genetic studies and concluded that the eastern wolf and red wolf are separate species from the gray wolf, having originated in North America 150,000–300,000 years ago from the same line as coyotes. The Chambers review concluded that the subspecific status of C. l. arctos is doubtful, as Arctic wolf populations do not possess unique haplotypes.[8] However, the Chambers review became controversial, forcing the USF&WS to commission a peer review of it, known as NCAES (2014).[43] This peer review concluded unanimously that the Chambers review "is not accepted as consensus scientific opinion or best available science..."

The taxonomy of wolves in the coastal rainforests of British Columbia and southeastern Alaska has also followed a variable path, regarding the putative Vancouver Island wolf (C. l. crassodon) and Alexander Archipelago wolf (C. l. ligoni), respectively. Based on skull morphometrics, C. l. ligoni was recognized by Goldman (1944), Hall (1981) and Pedersen (1982) as a distinct population possibly warranting subspecific classification; however, Nowak (1996) considered it to be an isolated population of C. l. nubilis.[44] From 2005 to 2014 several studies and the NCAES (2014) peer review have found the pacific coastal wolves to have a phenotypically distinct genotype.[45][46][47][48][49]

Domestication[edit]

33,000-year-old skull of a domesticated gray wolf from Siberia

The gray wolf was the first animal and only large carnivore to be domesticated by humans.[50] The origin of the domestic dog has been controversial, and subject to numerous studies, with genetic data and paleontological evidence contradicting each other on the date and location of the first domestication event. Proposed centers of dog origins from genetic data have included the Middle East[51] and East Asia 32,000 years ago,[52] though this is inconsistent with the paleontological record, as the oldest dog remains in those areas are no older than 13,000 years.[23] Much older remains were discovered in Europe and Russia,[53] with one specimen discovered in Goyet, Belgium having been estimated to be 31,700 years old,[54] and another found in the Altai Mountains being dated to be 33,000 years old.[55]

Several studies on gray wolf and dog mitochondrial genomes in 2013-14 showed that modern dog genomes don't match those of extant gray wolf populations, and likely arose from a now extinct lineage of prehistoric gray wolves in Europe, with any similarity with modern wolf populations reflecting historical admixture rather than recent divergence. The genetic data is consistent with the paleontological record, which shows that the oldest dog remains in Europe are 18,800 to 32,100 years old. The domestication event may have begun during the Last Glacial Maximum, prior to the Neolithic Revolution, when hunter-gatherers actively hunted Pleistocene megafauna. The morphology and DNA of the Goyet and Altai specimens didn't match those of modern dogs, thus indicating that they were the result of separate, ultimately aborted domestication episodes.[23][50] Further studies indicate that prehistoric Eurasian gray wolves underwent a threefold population bottleneck approximately 15,000-20,000 years ago, thus indicating that these gray wolves had substantially more genetic diversity for selection to act on than what is observed in modern wolf populations.[23]

Although the genetic divergence between gray wolves and dogs is only 1.8%, as opposed to over 4% between gray wolves, Ethiopian wolves and coyotes,[56] there are a number of diagnostic features to distinguish the two. The tympanic bullae are large, convex and almost spherical in gray wolves, while the bullae of dogs are smaller, compressed and slightly crumpled.[57] The teeth of gray wolves are also proportionately larger than those of dogs; the premolars and molars of wolves are much less crowded, and have more complex cusp patterns.[58] Dogs lack a functioning pre-caudal gland, and most enter estrus twice yearly, unlike gray wolves which only do so once a year.[59]

Hybridization with dogs and other Canis[edit]

Main article: Canid hybrids

Although dogs and gray wolves are genetically very close, and have shared vast portions of their ranges for millennia, the two generally do not voluntarily interbreed in the wild, though they can produce viable offspring, with all subsequent generations being fertile.[59] In North America, black colored wolves acquired their coloration from wolf-dog hybridization, which occurred 10,000–15,000 years ago.[60] Although wolf-dog hybridization in Europe has raised concern among conservation groups fearing for the gray wolf's purity, genetic tests show that introgression of dog genes into European gray wolf populations does not pose a significant threat. Also, as wolf and dog mating seasons do not fully coincide, the likelihood of wild wolves and dogs mating and producing surviving offspring is small.[61] Like pure wolves, hybrids breed once annually, though their mating season occurs three months earlier, with pups mostly being born in the winter period, thus lessening their chances of survival.[59] However, one genetic study undertaken in the Caucasus Mountains showed that as many as 10% of dogs in the area, including livestock guardian dogs, are first generation hybrids.[40] The captive breeding of wolf-dog hybrids has proliferated in the USA, with 300,000 such animals being present there.[59]

F1 hybrid coyote-gray wolf hybrid, conceived in captivity

The gray wolf has interbred extensively with the eastern wolf across northern Ontario, into Manitoba and Quebec, as well as into the western Great Lakes states of Minnesota, Wisconsin, and Michigan, producing a hybrid population termed Great Lakes boreal wolves.[62] The boreal wolf is 25% larger than a pure eastern wolf, and typically has a similarly colored gray-fawn coat but, unlike the eastern wolf, can also be black, cream, or white. It also specalises on larger prey such as moose and caribou rather than white-tailed deer. Unlike pure eastern wolves, Great Lakes boreal wolves primarily inhabit boreal rather than deciduous forests.[8]

Unlike the red and eastern wolf, the gray wolf does not readily interbreed with coyotes.[8] Nevertheless, coyote genetic markers have been found in some wild isolated gray wolf populations in the southern United States. Gray wolf Y-chromosomes have also been found in Texan coyote haplotypes.[63] In tests performed on a putative chupacabra carcass, mtDNA analysis showed that it was a coyote, though subsequent tests revealed that it was a coyote–gray wolf hybrid sired by a male Mexican gray wolf.[64] In 2013, a captive breeding experiment in Utah between gray wolves and western coyotes produced six hybrids through artificial insemination, making this the very first hybridization case between pure coyotes and northwestern gray wolves. At six months of age, the hybrids were closely monitored and were shown to display both physical and behaviourial characteristics from both species.[14]

Although hybridization between wolves and golden jackals has never been observed, evidence of such occurrences was discovered through mtDNA analysis on jackals in Senegal[15] and Bulgaria.[16] Although there is no genetic evidence of gray wolf-jackal hybridization in the Caucasus Mountains, there have been cases where otherwise genetically pure golden jackals have displayed remarkably gray wolf-like phenotypes, to the point of being mistaken for wolves by trained biologists.[40]

Physical description[edit]

Gray wolf skeleton, Abruzzo National Park, Italy

Anatomy and dimensions[edit]

See also: Dog anatomy
Gray wolf trotting, Denali National Park. The gray wolf generally places its hind paws in the tracks made by the front paws.[65]

Compared to its closest cousins (the coyote and golden jackal), the gray wolf is larger and heavier, with a broader snout, shorter ears, a shorter torso and longer tail.[11][15][57] It is a slender, powerfully built animal with a large, deeply descending ribcage, a sloping back and a heavily muscled neck.[11] The wolf's legs are moderately longer than those of other canids, which enables the animal to move swiftly, and allows it to overcome the deep snow that covers most of its geographical range.[65] Females tend to have narrower muzzles and foreheads, thinner necks, slightly shorter legs and less massive shoulders than males.[66] The gray wolf's head is large and heavy, with a wide forehead, strong jaws and a long, blunt muzzle. The ears are relatively small and triangular.[11] The teeth are heavy and large, being better suited to crushing bone than those of other extant canids, though not as specialised as those found in hyenas.[67][68] Its molars have a flat chewing surface, but not to the same extent as the coyote, whose diet contains more vegetable matter.[5] The gray wolf's jaws can exert a crushing pressure of perhaps 10,340 kPa (1,500 psi) compared to 5,200 kPa (750 psi) for a German shepherd. This force is sufficient to break open most bones.[69] The gray wolf usually carries its head at the same level as the back, raising it only when alert.[11] It usually travels at a loping pace, placing its paws one directly in front of the other. This gait can be maintained for hours at a rate of 8–9 km/h,[70] and allows the wolf to cover great distances. On bare paths, a wolf can quickly achieve speeds of 50–60 km/h. The gray wolf has a running gait of 55 to 70 km/h, can leap 5 metres horizontally in a single bound, and can maintain rapid pursuit for at least 20 minutes.[71]

Gray wolf and golden jackal exhibit at The Museum of Zoology, St. Petersburg. Note the wolf's larger size and broader muzzle.

The gray wolf is the largest extant member of the Canidae, excepting certain large breeds of domestic dog.[57] Gray wolf weight and size can vary greatly worldwide, tending to increase proportionally with latitude as predicted by Bergmann's Rule,[11] with the large wolves of Alaska and Canada sometimes weighing 3–6 times more than their Middle Eastern and South Asian cousins.[72] On average, adult wolves measure 105–160 cm (41–63 in) in length and 80–85 cm (32–34 in) in shoulder height.[11] The tail measures 29–50 cm (11–20 in) in length. The ears are 90–110 millimetres (3.5–4.3 in) in height, and the hind feet are 220–250 mm.[11] The skull averages 9–11 inches in length, and 5–6 inches wide.[73] Gray wolf weight varies geographically; on average, European wolves may weigh 38.5 kilograms (85 lb), North American wolves 36 kilograms (79 lb) and Indian and Arabian wolves 25 kilograms (55 lb).[74] Females in any given wolf population typically weigh 5–10 lbs less than males.[75] Wolves weighing over 54 kg (120 lbs) are uncommon, though exceptionally large individuals have been recorded in Alaska, Canada,[75] and the former Soviet Union.[11][76] The heaviest recorded gray wolf in North America was killed on 70 Mile River in east-central Alaska on July 12, 1939 and weighed 79.4 kilograms (175 lb),[77] while the heaviest recorded wolf in Eurasia was killed after World War II in Ukraine's Poltavskij Region, and weighed 86 kilograms (190 lb).[76]

Fur[edit]

Black and white-furred gray wolves, Pays de la Loire.

The gray wolf has very dense and fluffy winter fur, with short underfur and long, coarse guard hairs.[11] Most of the underfur and some of the guard hairs are shed in the spring and grow back in the autumn period.[74] The longest hairs occur on the back, particularly on the front quarters and neck. Especially long hairs are found on the shoulders, and almost form a crest on the upper part of the neck. The hairs on the cheeks are elongated and form tufts. The ears are covered in short hairs which strongly project from the fur. Short, elastic and closely adjacent hairs are present on the limbs from the elbows down to the calcaneal tendons.[11] The winter fur is highly resistant to cold; wolves in northern climates can rest comfortably in open areas at −40° by placing their muzzles between the rear legs and covering their faces with their tail. Wolf fur provides better insulation than dog fur, and does not collect ice when warm breath is condensed against it.[74] In warm climates, the fur is coarser and scarcer than in northern wolves.[11] Female wolves tend to have smoother furred limbs than males, and generally develop the smoothest overall coats as they age. Older wolves generally have more white hairs in the tip of the tail, along the nose and on the forehead. The winter fur is retained longest in lactating females, though with some hair loss around their nipples.[66] Hair length on the middle of the back is 60–70 mm. Hair length of the guard hairs on the shoulders generally does not exceed 90 mm, but can reach 110–130 mm.[11]

Coat color ranges from almost pure white through various shades of blond, cream, and ochre to grays, browns, and blacks,[78] with variation in fur color tending to increase in higher latitudes.[79] Differences in coat colour between sexes are largely absent,[11] though females may have redder tones.[80] Black coloured wolves (which occur through wolf-dog hybridisation) rarely occur in Eurasia, where interactions with domestic dogs have been reduced over the past thousand years due to the depletion of wild wolf populations. Black specimens are more common in North America, with about half the wolves in Yellowstone National Park being black.[60]

Behavior[edit]

See also: Dog behavior

Social and territorial behaviors[edit]

Gray wolf pack in Yellowstone National Park

The gray wolf is a social animal, whose basic social unit consists of a mated pair, accompanied by the pair's adult offspring.[d] The average pack consists of a family of 5–11 animals (1–2 adults, 3–6 juveniles and 1–3 yearlings),[11] or sometimes two or three such families,[5] with exceptionally large packs consisting of 42 wolves being known.[82] Triggers for dispersal include the onset of sexual maturity and competition within the pack for food.[83] In ideal conditions, the mated pair produces pups every year, with such offspring typically staying in the pack for 10–54 months before dispersing.[17] The distance travelled by dispersing wolves varies widely; some stay in the vicinity of the parental group, while other individuals may travel great distances of 390 km, 206 km, and 670 km from their natal packs.[84] A new pack is usually founded by an unrelated dispersing male and female, travelling together in search of an area devoid of other hostile packs.[85] Wolf packs rarely adopt other wolves into their fold, and typically kill them. In the rare cases where other wolves are adopted, the adoptee is almost invariably an immature animal (1–3 years of age) unlikely to compete for breeding rights with the mated pair. In some cases, a lone wolf is adopted into a pack to replace a deceased breeder.[82] During times of ungulate abundance (migration, calving etc.), different wolf packs may temporarily join forces.[11]

Wolves are highly territorial animals, and generally establish territories far larger than they require to survive in order to assure a steady supply of prey. Territory size depends largely on the amount of prey available and the age of the pack's pups, tending to increase in size in areas with low prey populations[86] or when the pups reach the age of 6 months, thus having the same nutritional needs as adults.[87] Wolf packs travel constantly in search of prey, covering roughly 9% of their territory per day (average 25 km/d or 15 mi/d). The core of their territory is on average 35 km2 (14 sq mi), in which they spend 50% of their time.[86] Prey density tends to be much higher in the territory's surrounding areas, though wolves tend to avoid hunting in the fringes of their range unless desperate, due to the possibility of fatal encounters with neighboring packs.[88] The smallest territory on record was held by a pack of six wolves in northeastern Minnesota, which occupied an estimated 33 km2 (13 sq mi), while the largest was held by an Alaskan pack of ten wolves encompassing a 6,272 km2 (2,422 sq mi) area.[87] Wolf packs are typically settled, and usually only leave their accustomed ranges during severe food shortages.[11]

Wolves defend their territories from other packs through a combination of scent marking, direct attacks and howling (see Communication). Scent marking is used for territorial advertisement, and involves urination, defecation and ground scratching.[89][90][91][92][93] Scent marks are generally left every 240 metres throughout the territory on regular travelways and junctions. Such markers can last for 2–3 weeks,[87] and are typically placed near rocks, boulders, trees or the skeletons of large animals.[11] Territorial fights are among the principal causes of wolf mortality, with one study concluding that 14–65% of wolf deaths in Minnesota and the Denali National Park and Preserve were due to predation by other wolves.[94]

Reproduction and development[edit]

Gray wolves mating, Tama zoo, Japan

The gray wolf is generally monogamous, with mated pairs usually remaining together for life, unless one of the pair dies. Upon the death of one mated wolf, pairs are quickly re-established. Since males often predominate in any given wolf population, unpaired females are a rarity.[11] If a dispersing male gray wolf is unable to establish a territory or find a mate, he mates with the daughters of already established breeding pairs from other packs. Such gray wolves are termed "Casanova wolves" and, unlike males from established packs, they do not form pair bonds with the females they mate with. Some gray wolf packs may have multiple breeding females this way, as is the case in Yellowstone National Park.[95] Gray wolves also practice alloparental care, in which a wolf pair may adopt the pup or pups of another. This might take place if the original parents die or are for some reason separated from them.[96] In addition to heterosexual behavior, homosexual behavior has been observed in gray wolves.[97] Male gray wolves often mount each other when the highest ranking female in the pack comes into heat.[98]

Gray wolf pups, Tiergarten Schönbrunn, Austria

The age of first breeding in gray wolves depends largely on environmental factors: when food is plentiful, or when wolf populations are heavily managed, wolves can rear pups at younger ages in order to better exploit abundant resources. This is further demonstrated by the fact that captive wolves have been known to breed as soon as they reach 9–10 months, while the youngest recorded breeding wolves in the wild were 2 years old. Females are capable of producing pups every year, with one litter annually being the average. Unlike the coyote, the gray wolf never reaches reproductive senescence.[99] Estrus typically occurs in late winter, with older, multiparous females entering estrus 2–3 weeks earlier than younger females.[11] During pregnancy, female wolves remain in a den located away from the peripheral zone of their territories, where violent encounters with other packs are more likely.[100] Old females usually whelp in the den of their previous litter, while younger females typically den near their birthplace. The gestation period lasts 62–75 days, with pups usually being born in the summer period.[11]

Wolves bear relatively large pups in small litters compared to other canid species.[101] The average litter consists of 5–6 pups,[102] with litter sizes tending to increase in areas where prey is abundant,[102] though exceptionally large litters of 14–17 pups occur only 1% of the time.[103] Pups are usually born in spring, coinciding with a corresponding increase in prey populations.[100] Pups are born blind and deaf, and are covered in short soft grayish-brown fur. They weigh 300–500 grams at birth, and begin to see after 9–12 days. The milk canines erupt after one month. Pups first leave the den after 3 weeks. At 1.5 months of age, they are agile enough to flee from danger. Mother wolves do not leave the den for the first few weeks, relying on the fathers to provide food for them and their young. Pups begin to eat solid food at the age of 3–4 weeks. Pups have a fast growth rate during their first four months of life: during this period, a pup's weight can increase nearly 30 times.[11][101] Wolf pups begin play fighting at the age of 3 weeks, though unlike young foxes and coyotes, their bites are inhibited. Actual fights to establish hierarchy usually occur at 5–8 weeks of age. This is in contrast to young foxes and coyotes, which may begin fighting even before the onset of play behavior.[12] By autumn, the pups are mature enough to accompany adults on hunts for large prey.[100]

Hunting and feeding behaviours[edit]

American Bison standing its ground, thereby increasing its chance of survival

Although social animals, single wolves or mated pairs typically have higher success rates in hunting than do large packs, with single wolves having occasionally been observed to kill large prey such as moose, bison and muskoxen unaided.[104] The gray wolf's sense of smell is relatively weakly developed when compared to that of some hunting dog breeds, being able to detect carrion upwind no farther than 2–3 km. Because of this, it rarely manages to capture hidden hares or birds, though it can easily follow fresh tracks. Its auditory perception is acute enough to be able to hear up to a frequency of 26 kHz,[105] which is sufficient to register the fall of leaves in the autumn period.[11] A gray wolf hunt can be divided into five stages:

  • Locating prey: The wolves travel in search of prey through their power of scent, chance encounter, and tracking. Wolves typically locate their prey by scent, though they must usually be directly downwind of it. When a breeze carrying the prey's scent is located, the wolves stand alert, and point their eyes, ears and nose towards their target. In open areas, wolves may precede the hunt with group ceremonies involving standing nose-to-nose and wagging their tails. Once concluded, the wolves head towards their prey.[106]
  • The stalk: The wolves attempt to conceal themselves as they approach.[107] As the gap between the wolves and their prey closes, the wolves quicken their pace, wag their tails, and peer intently, getting as close to their quarry as possible without making it flee.[108]
  • The encounter: Once the prey detects the wolves, it can either approach the wolves, stand its ground, or flee. Large prey, such as moose, elk, and muskoxen, usually stand their ground. Should this occur, the wolves hold back, as they require the stimulus of a running animal to proceed with an attack.[109] If the targeted animal stands its ground, the wolves either ignore it, or try to intimidate it into running.[104]
  • The rush: If the prey attempts to flee, the wolves immediately pursue it. This is the most critical stage of the hunt, as wolves may never catch up with prey running at top speed.[110] If their prey is travelling in a group, the wolves either attempt to break up the herd, or isolate one or two animals from it.[107]
  • The chase: A continuation of the rush, the wolves attempt to catch up with their prey and kill it.[111] When chasing small prey, wolves attempt to catch up with their prey as soon as possible, while with larger animals, the chase is prolonged, in order to wear the selected prey out. Wolves usually give up chases after 1–2 km (0.62–1.3 mi), though one wolf was recorded to chase a deer for 21 km (13 mi).[104] Both Russian and North American wolves have been observed to drive prey onto crusted ice, precipices, ravines, slopes and steep banks to slow them down.[112]
Killing a caribou in typical fashion: biting the hindquarters (1909 drawing by A. R. Harding)

The actual killing method varies according to prey species. With large prey, mature wolves usually avoid attacking frontally, instead focusing on the rear and sides of the animal. Large prey, such as moose, is killed by biting large chunks of flesh from the soft perineum area, causing massive blood loss. Such bites can cause wounds 10–15 cm in length, with three such bites to the perineum usually being sufficient to bring down a large deer in optimum health.[112] With medium-sized prey such as roe deer or sheep, wolves kill by biting the throat, severing nerve tracks and the carotid artery, thus causing the animal to die within a few seconds to a minute. With small, mouse-like prey, wolves leap in a high arc and imobilize it with their forepaws.[113] When prey is vulnerable and abundant, wolves may occasionally surplus kill. Such instances are common in domestic animals, but rare in the wild. In the wild, surplus killing primarily occurs during late winter or spring, when snow is unusually deep (thus impeding the movements of prey)[114] or during the denning period, when wolves require a ready supply of meat when denbound.[115] Medium-sized prey are especially vulnerable to surplus killing, as the swift throat-biting method by which they are killed allows wolves to quickly kill one animal and move on to another.[113]

Once prey is brought down, wolves begin to feed excitedly, ripping and tugging at the carcass in all directions, and bolting down large chunks of it.[116] The breeding pair typically monopolizes food in order to continue producing pups. When food is scarce, this is done at the expense of other family members, especially non-pups.[117] The breeding pair typically eats first, though as it is they who usually work the hardest in killing prey, they may rest after a long hunt and allow the rest of the family to eat unmolested. Once the breeding pair has finished eating, the rest of the family tears off pieces of the carcass and transport them to secluded areas where they can eat in peace. Wolves typically commence feeding by consuming the larger internal organs of their prey, such as the heart, liver, lungs and stomach lining. The kidneys and spleen are eaten once they are exposed, followed by the muscles.[118] A single wolf can eat 15–19% of its body weight in a single feeding.[119]

Communication[edit]

Visual[edit]

Facial expressions (Konrad Lorenz, 1952). bottom to top: increasing fear (ears back); left to right: increasing aggression (snarl); top right: maximum of both.

The gray wolf's expressive behavior is more complex than that of the coyote and golden jackal, as necessitated by its group living and hunting habits. While less gregarious canids generally possess simple repertoires of visual signals, wolves have more varied signals which subtly inter grade in intensity.[12][13] When neutral, the legs are not stiffened, the tail hangs down loosely, the face is smooth, the lips untensed, and the ears point in no particular direction.[120] Postural communication in wolves consists of a variety of facial expressions, tail positions and piloerection.[105] Aggressive, or self-assertive wolves are characterized by their slow and deliberate movements, high body posture and raised hackles, while submissive ones carry their bodies low, sleeken their fur and lower their ears and tail.[121] When a breeding male encounters a subordinate family member, it may stare at it, standing erect and still with the tails horizontal to its spine.[122] Two forms of submissive behavior are recognized: passive and active. Passive submission usually occurs as a reaction to the approach of a dominant animal, and consists of the submissive wolf lying partly on its back and allowing the dominant wolf to sniff its anogenital area. Active submission occurs often as a form of greeting, and involves the submissive wolf approaching another in a low posture, and licking the other wolf's face.[123] When wolves are together, they commonly indulge in behaviors such as nose pushing, jaw wrestling, cheek rubbing and facial licking. The mouthing of each other's muzzles is a friendly gesture, while clamping on the muzzle with bared teeth is a dominance display.[124]

Expressive characteristics of visual features
used during social interactions in wolves[121]
FeatureAggressiveFearful
EyesDirect stare
Open wide
Looking away
Closed to slits
EarsErect and forwardFlattened and turned down to side
LipsHorizontal contraction
("agonistic pucker")
Horizontal retraction ("submissive grin")
MouthOpenedClosed
TeethCanines baredCanines covered
TongueRetractedExtended ("lick intention")
NoseShortened (skin folded)Lengthened (skin smoothed)
ForeheadContracted (bulging over eyes)Stretched (smoothed)
HeadHeld highLowered
NeckArchedExtended
HairErect (bristled)Sleeked
BodyErect, tallCrouched, low
TailHeld high
Quivering
Tucked under body
Wagging

Auditory[edit]

Howling gray wolf, UK Wolf Conservation Trust

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Gray wolves howl to assemble the pack (usually before and after hunts), to pass on an alarm (particularly at a den site), to locate each other during a storm or unfamiliar territory and to communicate across great distances.[125] Wolf howls can under certain conditions be heard over areas of up to 130 km2 (50 sq mi).[5] Wolf howls are generally indistinguishable from those of large dogs.[126] Male wolves give voice through an octave, passing to a deep bass with a stress on "O", while females produce a modulated nasal baritone with stress on "U". Pups almost never howl, while yearling wolves produce howls ending in a series of dog-like yelps.[11] Howling consists of a fundamental frequency which may lie between 150 and 780 Hz, and consists of up to 12 harmonically related overtones. The pitch usually remains constant or varies smoothly, and may change direction as many as four or five times.[57] Howls used for calling pack mates to a kill are long, smooth sounds similar to the beginning of the cry of a horned owl. When pursuing prey, they emit a higher pitched howl, vibrating on two notes. When closing in on their prey, they emit a combination of a short bark and a howl.[126] When howling together, wolves harmonize rather than chorus on the same note, thus creating the illusion of there being more wolves than there actually are.[125] Lone wolves typically avoid howling in areas where other packs are present.[127] Wolves from different geographic locations may howl in different fashions: the howls of European wolves are much more protracted and melodious than those of North American wolves, whose howls are louder and have a stronger emphasis on the first syllable. The two are however mutually intelligible, as North American wolves have been recorded to respond to European-style howls made by biologists.[128]

Other vocalisations of wolves are usually divided into three categories: growls, barks and whines.[129] Barking has a fundamental frequency between 320–904 Hz,[57] and is usually emitted by startled wolves. Wolves do not bark as loudly or continuously as dogs do, but bark a few times and retreat from perceived danger.[129] Growling has a fundamental frequency of 380–450 Hz,[57] and is usually emitted during food challenges. Pups commonly growl when playing. One variation of the howl is accompanied by a high pitched whine, which precedes a lunging attack.[125] Whining is associated with situations of anxiety, curiosity, inquiry and intimacy such as greeting, feeding pups and playing.[129]

Olfactory[edit]

Gray wolf marking its territory, Kolmården Zoo. Raised leg urination (RLU) is considered the most important form of scent communication in the wolf, making up 60–80% of all scent marks observed.[130]

Olfaction is probably the wolf's most acute sense, and plays a fundamental role in communication. The wolf has a large number of apocrine sweat glands on the face, lips, back, and between the toes. The odor produced by these glands varies according to the individual wolf's microflora and diet, giving each a distinct "odor fingerprint". A combination of apocrine and eccrine sweat glands on the feet allows the wolf to deposit its scent whilst scratching the ground, which usually occurs after urine marking and defecation during the breeding season. The follicles present on the guard hairs from the wolf's back have clusters of apocrine and sebaceous glands at their bases. As the skin on the back is usually folded, this provides a microclimate for bacterial propagation around the glands. During piloerection, the guard hairs on the back are raised and the skin folds spread, thus releasing scent.[131] The pre-caudal scent glands may play a role in expressing aggression, as combative wolves raise the base of their tails whilst drooping the tip, thus positioning the scent glands at the highest point.[126] The wolf possesses a pair of anal sacs beneath the rectum, which contain both apocrine and sebaceous glands. The components of anal sac secretions vary according to season and gender, thus indicating that the secretions provide information related to gender and reproductive state. The secretions of the preputial glands may advertise hormonal condition or social position, as dominant wolves have been observed to stand over subordinates, apparently presenting the genital area for investigation,[131] which may include genital licking.[132] During the breeding season, female wolves secrete substances from the vagina which communicate the females' reproductive state, and can be detected by males from long distances. Urine marking is the best-studied means of olfactory communication in wolves. Its exact function is debated, though most researchers agree that its primary purpose is to establish boundaries. Wolves urine mark more frequently and vigorously in unfamiliar areas, or areas of intrusion, where the scent of other wolves or canids is present. So-called raised leg urination (RLU) is more common in male wolves than in females, and may serve the purpose of maximizing the possibility of detection by conspecifics, as well as reflect the height of the marking wolf. Only dominant wolves typically use RLU, with subordinate males continuing to use the juvenile standing posture throughout adulthood.[131]

Ecology[edit]

Habitat[edit]

The gray wolf is a habitat generalist, and can occur in deserts, grasslands, forests and arctic tundras. Habitat use by gray wolves is strongly corrolated with the abundance of prey, snow conditions, absence or low livestock densities, road densities, human presence and topography.[5] In cold climates, the gray wolf can reduce the flow of blood near its skin to conserve body heat. The warmth of the footpads is regulated independently of the rest of the body, and is maintained at just above tissue-freezing point where the pads come in contact with ice and snow.[133] Gray wolves use different places for their diurnal rest: places with cover are preferred during cold, damp and windy weather, while wolves in dry, calm and warm weather readily rest in the open. During the autumn-spring period, when wolves are more active, they willingly lie out in the open, whatever their location. Actual dens are usually constructed for pups during the summer period. When building dens, females make use of natural shelters such as fissures in rocks, cliffs overhanging riverbanks and holes thickly covered by vegetation. Sometimes, the den is the appropriated burrow of smaller animals such as foxes, badgers or marmots. An appropriated den is often widened and partly remade. On rare occasions, female wolves dig burrows themselves, which are usually small and short with 1–3 openings. The den is usually constructed not more than 500 metres away from a water source,[11] and typically faces southwards, thus ensuring enough sunlight exposure, keeping the denning area relatively snow free.[5] Resting places, play areas for the pups and food remains are commonly found around wolf dens. The odour of urine and rotting food emanating from the denning area often attracts scavenging birds such as magpies and ravens. As there are few convenient places for burrows, wolf dens are usually occupied by animals of the same family. Though they mostly avoid areas within human sight, wolves have been known to nest near domiciles, paved roads and railways.[11]

Diet[edit]

Gray wolf carrying caribou hindquarter, Denali National Park

The gray wolf generally specializes in vulnerable individuals of large prey.[5] In Eurasia, many gray wolf populations are forced to subsist largely on livestock and garbage in areas with dense human activity, though wild ungulates such as moose, red deer, roe deer and wild boar are still the most important food sources in Russia and the more mountainous regions of Eastern Europe. Other prey species include reindeer, argali, mouflon, wisent, saiga, ibex, chamois, wild goats, fallow deer and musk deer.[134] The prey animals of North American wolves have largely continued to occupy suitable habitats with low human density, and cases of wolves subsisting largely on garbage or livestock are exceptional. Animals preferred as prey by North American wolves include moose, white-tailed deer, elk, mule deer, bighorn sheep, Dall's sheep, American bison, muskox and caribou.[135]

Although wolves primarily feed on medium to large sized ungulates, they are not fussy eaters. Smaller sized animals that may supplement the diet of wolves include marmots, hares, badgers, foxes, weasels, ground squirrels, mice, hamsters, voles and other rodents, as well as insectivores. They frequently eat waterfowl and their eggs. When such foods are insufficient, they prey on lizards, snakes, frogs, rarely toads and large insects as available. In times of scarcity, wolves readily eat carrion, visiting cattle burial grounds and slaughter houses.[11] Cannibalism is not uncommon in wolves: during harsh winters, packs often attack weak or injured wolves, and may eat the bodies of dead pack members.[11][136][137] Wolf packs in Astrakhan hunt Caspian seals on the Caspian Sea coastline[138] and some wolf packs in Alaska and Western Canada have been observed to feed on salmon.[139] Humans are rarely, but occasionally preyed upon.[11][140] Other primates occasionally taken by wolves include grey langurs in Nepal[141] and hamadryas baboons in Saudi Arabia.[142]

Wolves supplement their diet with fruit and vegetable matter. They willingly eat the berries of mountain ash, lily of the valley, bilberries, blueberries and cowberry. Other fruits include nightshade, apples and pears. They readily visit melon fields during the summer months.[11] A well-fed wolf stores fat under the skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter.[119] Digestion only takes a few hours, thus wolves can feed several times in one day, making quick use of large quantities of meat.[143]

Enemies and competitors[edit]

Gray wolves confronting coyotes over pronghorn carcass (1919), Louis Agassiz Fuertes.

Gray wolves typically dominate other canid species in areas where they both occur. In North America, incidents of gray wolves killing coyotes are common, particularly in winter, when coyotes feed on wolf kills. Wolves may attack coyote den sites, digging out and killing their pups, though rarely eating them. There are no records of coyotes killing wolves, though coyotes may chase wolves if they outnumber them.[144][145] Near identical interactions have been observed in Eurasia and Africa between gray wolves and golden jackals, with the latter's numbers being comparatively small in areas with high wolf densities.[11][15][144][146] Gray wolves are the most important predator of raccoon dogs, killing large numbers of them in the spring and summer periods.[11] Wolves also kill red, arctic and corsac foxes, usually in disputes over carcasses, sometimes eating them.[11][147] In Asia, they may compete with dholes.[11]

Brown bears typically dominate wolf packs in disputes over carcasses, while wolf packs mostly prevail against bears when defending their den sites. Both species kill each other's young. Wolves eat the brown bears they kill, while brown bears seem to only eat young wolves.[148] Wolf interactions with American black bears 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 on numerous occasions to actively seek out black bears in their dens and kill them without eating them. Unlike brown bears, black bears frequently lose against wolves in disputes over kills.[149] While encounters with brown and black bears appear to be common, polar bears are rarely encountered by wolves, though there are two records of wolf packs killing polar bear cubs.[150] Wolves also kill the cubs of Asian black bears.[11]

Wolves may encounter striped hyenas in Israel and Central Asia, usually in disputes over carcasses. Striped hyenas feed extensively on wolf-killed carcasses in areas where the two species interact. One-to-one, hyenas dominate wolves, but wolf packs can drive off single hyenas.[151]

Large wolf populations limit the numbers of small to medium sized felines. Wolves encounter cougars along portions of the Rocky Mountains and adjacent mountain ranges. Wolves and cougars typically avoid encountering each other by hunting on different elevations. In winter however, when snow accumulation forces their prey into valleys, interactions between the two species become more likely. Wolves in packs usually dominate cougars and can steal kills. They have been reported killing mothers and their kittens.[152][153] Wolves hunt steppe cats, and may pose a threat to snow leopards.[154] Wolves may also reduce Eurasian lynx populations. Wolves may kill lynxes by running them down, or killing them before they can escape into trees.[155] Similar reports of encounters between wolves and bobcats have been documented.[156]

Leftovers of wolf kills are sometimes scavenged by wolverines. Wolverines usually wait until the wolves are done feeding, but have been known to drive away wolves from kills. However, there have been confirmed reports of wolf packs killing wolverines.[157]

Other than humans, tigers appear to be the only serious predators of wolves.[11][18][19][20][154] Wolf and tiger interactions are well documented in Sikhote-Alin, where 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 tiger numbers. Proven cases of tigers killing wolves are rare and attacks appear to be competitive rather than predatory in nature, with at least four proven records of tigers killing wolves without consuming them.[158]

Range and conservation[edit]

Present and past distribution of the gray wolf (not including the newly discovered wolf populations in Ethiopia, Egypt, Senegal, Mali and Algeria)

Europe[edit]

Decline[edit]

Polychrome cave painting of a gray wolf, Font-de-Gaume, France.

The extermination of Northern Europe's wolves first became an organized effort during the Middle Ages, and continued until the late 1800s. In England, wolf persecution was enforced by legislation, and the last wolf was killed in the early sixteenth century during the reign of Henry VII. Wolves lasted longer in Scotland, where they sheltered in vast tracts of forest, which were subsequently burned down. Wolves managed to survive in the forests of Braemar and Sutherland until 1684. The extirpation of wolves in Ireland followed a similar course, with the last wolf believed to have been killed in 1786.[159] A wolf bounty was introduced in Sweden in 1647, after the extermination of moose and reindeer forced wolves to feed on livestock. The Sami extirpated wolves in northern Sweden in organized drives. By 1960, few wolves remained in Sweden, due to the use of snowmobiles in hunting them, with the last specimen being killed in 1966. The gray wolf was exterminated in Denmark in 1772 and Norway's last wolf was killed in 1973. The species was decimated in 20th century Finland, despite regular dispersals from Russia. The gray wolf was only present in the eastern and northern parts of Finland by 1900, though its numbers increased after World War II.[160]

In Central Europe, wolves were dramatically reduced in number during the early nineteenth century, due to organized hunts and reductions in ungulate populations. In Bavaria, the last wolf was killed in 1847, and had disappeared from the Rhine regions by 1899[160] and largely disappeared in Switzerland before the end of the nineteenth century. In 1934, Nazi Germany became the first state in modern history to place the wolf under protection, though the species was already extirpated in Germany at this point.[161] The last free-living wolf to be killed on the soil of present-day Germany before 1945 was the so-called "Tiger of Sabrodt", which was shot near Hoyerswerda, Lusatia (then Lower Silesia) in 1904. Today, wolves have returned to the area.[162] Wolf hunting in France was first institutionalized by Charlemagne between 800–813, when he established the louveterie, a special corps of wolf hunters. The louveterie was abolished after the French Revolution in 1789, but was re-established in 1814. In 1883, up to 1,386 wolves were killed, with many more by poison.[160]

Last wolf to be killed in central Finland (Karstula, 1911).

In Eastern Europe, wolves were never fully exterminated, due to the area's contiguity with Asia and its large forested areas. However, Eastern European wolf populations were reduced to very low numbers by the late nineteenth century. Wolves were extirpated in Slovakia during the first decade of the twentieth century and, by the mid-twentieth century, could only be found in a few forested areas in eastern Poland. Wolves in the eastern Balkans benefitted from the region's contiguity with the former Soviet Union and large areas of plains, mountains and farmlands. Wolves in Hungary occurred in only half the country around the start of the 20th century, and were largely restricted to the Carpathian Basin. Wolf populations in Romania remained largely substantial, with an average of 2,800 wolves being killed annually out of a population of 4,600 from 1955–1965. An all time low was reached in 1967, when the population was reduced to 1,550 animals. The extermination of wolves in Bulgaria was relatively recent, as a previous population of about 1,000 animals in 1955 was reduced to about 100–200 in 1964. In Greece, the species disappeared from the southern Peloponnese in 1930. Despite periods of intense hunting during the eighteenth century, wolves never disappeared in the western Balkans, from Albania to the former Yugoslavia. Organized persecution of wolves began in Yugoslavia in 1923, with the setting up of the Wolf Extermination Committee (WEC) in Kocevje, Slovenia. The WEC was successful in reducing wolf numbers in the Dinaric Alps.[160]

In Southern Europe, wolf extermination was not as complete as in Northern Europe, due to greater cultural tolerance of the species. Wolf populations only began declining in the Iberian Peninsula in the early 19th-century, and was reduced by a half of its original size by 1900. Wolf bounties were regularly paid in Italy as late as 1950. Wolves were exterminated in the alps by 1800, and numbered only 100 by 1973, inhabiting only 3–5% of their former Italian range.[160]

Recovery[edit]

Gray wolves in Bardu, Troms, Norway

The recovery of European wolf populations began after the 1950s, when traditional pastoral and rural economies declined and thus removed the need to heavily persecute wolves. By the 1980s, small and isolated wolf populations expanded in the wake of decreased human density in rural areas and the recovery of wild prey populations.[163]

The gray wolf has been fully protected in Italy since 1976, and now holds a population of 450–500 animals, which has an annual increase of about 60%. Italian wolves entered France's Mercantour National Park in 1993, and at least fifty wolves were discovered in the western Alps in 2000. By 2013 the 250 wolves in the Western Alps imposed a significant burden on traditional sheep and goat husbandry with a loss of over 5,000 animals in 2012.[164] There are approximately 2000 wolves inhabiting the Iberian Peninsula, of which 150 reside in northeastern Portugal. In Spain, the species occurs in Galicia, Leon, and Asturia. Although hundreds of Iberian wolves are illegally killed annually, the population has expanded south across the river Duero and east to the Asturias and Pyrenees Mountains.[163]

In 1978, wolves began recolonising central Sweden after a twelve-year absence, and have since expanded into southern Norway. As of 2005, the total number of Swedish and Norwegian wolves is estimated to be at least one hundred, including eleven breeding pairs. The gray wolf is fully protected in Sweden and partially controlled in Norway. The Scandinavian wolf populations owe their continued existence to neighbouring Finland's contiguity with the Republic of Karelia, which houses a large population of wolves. Wolves in Finland are protected only in the southern third of the country, and can be hunted in other areas during specific seasons,[163] though poaching remains common, with 90% of young wolf deaths being due to human predation, and the number of wolves killed exceeds the number of hunting licenses, in some areas by a factor of two. Furthermore, the decline in the moose populations has reduced the wolf's food supply.[165][166] Since 2011, Netherlands, Belgium and Denmark have also reported wolf sightings presumably by natural migration from adjacent countries.[167][168]

Wolf populations in Poland have increased to about 800–900 individuals since being classified as a game species in 1976. Poland plays a fundamental role in providing routes of expansion into neighbouring Central European countries. In the east, its range overlaps with populations in Lithuania, Belorussia, Ukraine, and Slovakia. A population in western Poland expanded into eastern Germany and in 2000 the first cubs were born on German territory.[169] In 2012, an estimated 14 wolf packs were living in Germany (mostly in the east) and a pack with cubs has been sighted within 15 miles of Berlin.[170] The gray wolf is protected in Slovakia, though an exception is made for wolves killing livestock. A few Slovakian wolves disperse into the Czech Republic, where they are afforded full protection. Wolves in Slovakia, Ukraine and Croatia may disperse into Hungary, where the lack of cover hinders the buildup of an autonomous population. Although wolves have special status in Hungary, they may be hunted with a year-round permit if they cause problems.[163]

Romania has a large population of wolves, numbering 2500 animals. The wolf has been a protected animal in Romania since 1996, although the law is not enforced. The number of wolves in Albania and Macedonia is largely unknown, despite the importance the two countries have in linking wolf populations from Greece to those of Bosnia and Herzegovina and Croatia. Although protected, many wolves are illegally killed in Greece annually, and their future is uncertain. Wolf numbers have declined in Bosnia and Herzegovina since 1986, while the species is fully protected in neighbouring Croatia and Slovenia.[163]

Asia[edit]

Historical range and decline[edit]

Monument to the last Japanese gray wolf, Nara Prefecture, Japan

During the 19th century, gray wolves were widespread in many parts of the Holy Land east and west of the Jordan River. However, they decreased considerably in number between 1964 and 1980, largely due to persecution by farmers.[171] The species was not considered common in northern and central Saudi Arabia during the 19th century, with most early publications involving animals either from southwestern Asir, northern rocky areas bordering Jodan, or areas surrounding Riyadh.[172]

The gray wolf's range in the Soviet Union encompassed nearly the entire territory of the country, being absent only on the Solovetsky Islands, Franz-Josef Land, Severnaya Zemlya, and the Karagin, Commander and Shantar Islands. The species was exterminated twice in Crimea, once after the Russian Civil War, and again after World War II.[11] Following the two world wars, Soviet wolf populations peaked twice. 30,000 wolves were harvested annually out of a population of 200,000 during the 1940s, with 40,000–50,000 harvested during peak years. Soviet wolf populations reached a low around 1970, disappearing over much of European Russia. The population increased again by 1980 to about 75,000, with 32,000 being killed in 1979.[173] Wolf populations in northern inner Mongolia declined during the 1940s, primarily due to poaching of gazelles, the wolf's main prey.[174] In British-ruled India, wolves were heavily persecuted due to their attacks on sheep, goats and children. In 1876, 2,825 wolves were bountied in the North-Western Provinces (NWP) and Bihar. By the 1920s, wolf extermination remained a priority in the NWP and Awadh. Overall, over 100,000 wolves were killed for bounties in British India between 1871 and 1916.[175]

Wolves in Japan became extinct during the Meiji restoration period, an extermination known as ōkami no kujo. The wolf was deemed a threat to ranching which the Meiji government promoted at the time, and targeted via a bounty system and a direct chemical extermination campaign inspired by the similar contemporary American campaign. The last Japanese wolf was a male killed on the 23 January 1905 near Washikaguchi (now called Higashi Yoshiro).[176]

Modern range[edit]

Gray wolf near Ardahan, Turkey. Although Turkish wolves have no legal protection, they may number about 7,000 individuals.[177]

There is little reliable data on the status of wolves in the Middle East, save for those in Israel and Saudi Arabia, though their numbers appear to be stable, and are likely to remain so. Israel's conservation policies and effective law enforcement maintain a moderately sized wolf population, which radiates into neighbouring countries, while Saudi Arabia has vast tracts of desert, where about 300–600 wolves live undisturbed.[178] The wolf survives throughout most of its historical range in Saudi Arabia, probably due to a lack of pastoralism and abundant human waste.[172] Turkey may play an important role in maintaining wolves in the region, due to its contiguity with Central Asia. The mountains of Turkey have served as a refuge for the few wolves remaining in Syria. A small wolf population occurs in the Golan Heights, and is well protected by the military activities there. Wolves living in the southern Negev desert are contiguous with populations living in the Egyptian Sinai and Jordan. Throughout the Middle East, the species is only protected in Israel. Elsewhere, it can be hunted year-round by Bedouins.[178]

Little is known of current wolf populations in Iran, which once occurred throughout the country in low densities during the mid-1970s. The northern regions of Afghanistan and Pakistan are important strongholds for the wolf. It has been estimated that there are about 300 wolves in approximately 60,000 km2 (23,000 sq mi) of Jammu and Kashmir in northern India, and 50 more in Himachal Pradesh. Overall, India supports about 800-3,000 wolves, scattered among several remnant populations. Although protected since 1972, Indian wolves are classed as Endangered, with many populations lingering in low numbers or living in areas increasingly used by humans. Although present in Nepal and Bhutan, there is no information of wolves occurring there.[173]

Wolf populations throughout Northern and Central Asia are largely unknown, but are estimated in the hundreds of thousands based on annual harvests. Since the fall of the Soviet Union, continent-wide extermination of wolves has ceased, and wolf populations have increased to about 25,000–30,000 animals throughout the former Soviet Union. In China and Mongolia, wolves are only protected in reserves. Mongolian populations have been estimated at 10,000–30,000, while the status of wolves in China is more fragmentary. The north has a declining population of an estimated 400 wolves, while Xinjiang and Tibet hold about 10,000 and 2,000 respectively.[179]

North America[edit]

A 100 lb gray wolf killed in Montana, 1928. Prior to their extirpation around 1930, Montana's wolves could be very large. Wolves recolonized the state from Canada beginning in the 1970s. Courtesy USFWS.

Historical range and decline[edit]

Originally, the gray wolf occupied all of North America north of about 20°N. It occurred all over the mainland, save for the southeastern USA, California west of the Sierra Nevada, and the tropical and subtropical areas of Mexico. Large continental islands occupied by wolves included Newfoundland, Vancouver Island, southeastern Alaskan islands, and throughout the Arctic Archipelago and Greenland.[5]

The decline of North American wolf populations coincided with increasing human populations and the expansion of agriculture. By the start of the 20th century, the species had almost disappeared from the eastern USA, excepting some areas of the Appalachians and the northwestern Great Lakes Region. In Canada, the gray wolf was extirpated in New Brunswick and Nova Scotia between 1870 and 1921, and in Newfoundland around 1911. It vanished from the southern regions of Quebec and Ontario between 1850 and 1900. The gray wolf's decline in the prairies began with the extermination of the American bison and other ungulates in the 1860s–70s. From 1900–1930, the gray wolf was virtually eliminated from the western USA and adjoining parts of Canada, due to intensive predator control programs aimed at eradicating the species. The gray wolf was exterminated by federal and state governments from all of the USA by 1960, except in Alaska and northern Minnesota. The decline in North American wolf populations was reversed from the 1930s to the early 1950s, particularly in southwestern Canada, due to expanding ungulate populations resulting from improved regulation of big game hunting. This increase triggered a resumption of wolf control in western and northern Canada. Thousands of wolves were killed from the early 1950s to the early 1960s, mostly due to poisoning. This campaign was halted and wolf populations increased again by the mid-1970s.[5]

Modern range[edit]

The species' modern range in North America is mostly confined to Alaska and Canada, with populations also occurring in northern Minnesota, northern Wisconsin and Michigan's Upper Peninsula, and portions of Washington, Idaho, northern Oregon, and Montana. A functional wolf population should exist in California by 2024 according to estimates by state wildlife officials.[180] Canadian wolves began to naturally re-colonize northern Montana around Glacier National Park in 1979, and the first wolf den in the western U.S. in over half a century was documented there in 1986.[181] The wolf population in northwest Montana initially grew as a result of natural reproduction and dispersal to about 48 wolves by the end of 1994.[182] From 1995-1996, wolves from Alberta and British Columbia were relocated to Yellowstone National Park and Idaho. In addition, the Mexican wolf (Canis lupus baileyi) was reintroduced to Arizona and New Mexico in 1998. The gray wolf is found in approximately 80% of its historical range in Canada, thus making it an important stronghold for the species.[5]

Canada is home to about 52,000–60,000 wolves, whose legal status varies according to province and territory. First Nations residents may hunt wolves without restriction, and some provinces require licenses for residents to hunt wolves while others do not. In Alberta, wolves on private land may be baited and hunted by the landowner without requiring a license, and in some areas, wolf hunting bounty programs exist.[183][184] Large-scale wolf population control through poisoning, trapping and aerial hunting is also presently conducted by government-mandated programs in order to support populations of endangered prey species such as Woodland caribou.[185]

In Alaska, the gray wolf population is estimated at 6,000–7,000, and can be legally harvested during hunting and trapping seasons, with bag limits and other restrictions. As of 2002, there are 250 wolves in 28 packs in Yellowstone, and 260 wolves in 25 packs in Idaho. The gray wolf received Endangered Species Act (ESA) protection in Minnesota, Wisconsin, and Michigan in 1974, and was re-classed from Endangered to Threatened in 2003. Reintroduced Mexican wolves in Arizona and New Mexico are protected under the ESA and, as of late 2002, number 28 individuals in eight packs.[186] A female wolf shot in 2013 in Hart County, Kentucky by a hunter was the first gray wolf seen in Kentucky in modern times. DNA analysis by Fish and Wildlife laboratories showed genetic characteristics similar to those of wolves in the Great Lakes Region.[187]

Africa[edit]

Up until 2011, it was debated whether or not the gray wolf occurred in Africa. Aristotle wrote of wolves living in Egypt, mentioning that they were smaller than the Greek kind. Georg Ebers wrote of the wolf being among the sacred animals of Egypt, describing it as a "smaller variety" of wolf to those of Europe, and noting how the name Lykopolis, the Ancient Egyptian city dedicated to Anubis, means "city of the wolf". Zoologist Ernst Schwarz classified North Africa's wolf-like canid as a subspecies of golden jackal, and was subsequently criticised for having overlooked its morphological affinity to the gray wolf.[39][188] In December 2002, a canid was sighted in Eritrea's Danakil Desert, whose appearance didn't correspond to that of the golden jackal or the six other recognised species of the area, but strongly resembled that of the gray wolf.[189] That the canid was a gray wolf was proven in 2011, when the base pairs of the mtDNA of samples taken from the Ethiopian Highlands were analyzed and compared with those of other wolves and wolf-like canids. The results suggested that African wolves inhabit at least two places in Ethiopia, approximately 2,500 km southeast of Egypt.[39] A further study confirmed the presence of gray wolves in Algeria, Mali and Senegal.[15]

Diseases and parasites[edit]

Viral and bacterial infections[edit]

Viral diseases carried by wolves include rabies, canine distemper, canine parvovirus, infectious canine hepatitis, papillomatosis, and canine coronavirus.[190] Wolves are a major host for rabies in Russia, Iran, Afghanistan, Iraq and India.[191] In wolves, the incubation period is 8–21 days, and results in the host becoming agitated, deserting its pack, and travelling up to 80 km a day, thus increasing the risk of infecting other wolves. Infected wolves do not show any fear of humans, with most documented wolf attacks on people being attributed to rabid animals. Although canine distemper is lethal in dogs, it has not been recorded to kill wolves, except in Canada and Alaska. The canine parvovirus, which causes death by dehydration, electrolyte imbalance, and endotoxic shock or sepsis, is largely surviveable in wolves, but can be lethal to pups. Wolves may catch infectious canine hepatitis from dogs, though there are no records of wolves dying from it. Papillomatosis has been recorded only once in wolves, and likely doesn't cause serious illness or death, though it may alter feeding behaviors. The canine coronavirus has been recorded in Alaskan wolves, with infections being most prevalent in winter months.[190]

Bacterial diseases carried by wolves include brucellosis, lyme disease, leptospirosis, tularemia, bovine tuberculosis,[192] listeriosis, anthrax and foot and mouth disease.[191] Wolves can catch Brucella suis from wild and domestic reindeer. While adult wolves tend not to show any clinical signs, it can severely weaken the pups of infected females. Although lyme disease can debilitate individual wolves, it does not appear to have any significant effect on wolf populations. Leptospirosis can be contracted through contact with infected prey or urine, and can cause fever, anorexia, vomiting, anemia, hematuria, icterus, and death. Wolves living near farms are more vulnerable to the disease than those living in the wilderness, probably due to prolonged contact with infected domestic animal waste. Wolves may catch tularemia from lagomorph prey, though its effect on wolves is unknown. Although bovine tuberculosis is not considered a major threat to wolves, it has been recorded to have once killed two wolf pups in Canada.[192]

Parasitic infections[edit]

Wolves carry ectoparasites and endoparasites, with wolves in the former Soviet Union having been recorded to carry at least 50 species.[191] Most of these parasites infect wolves without adverse effects, though the effects may become more serious in sick or malnourished specimens.[193] Parasitic infection in wolves is of particular concern to people, as wolves can spread them to dogs, which in turn can carry the parasites to humans. In areas where wolves inhabit pastoral areas, the parasites can be spread to livestock.[191]

Wolves are often infested with a variety of arthropod exoparasites, including fleas, ticks, lice, and mites. The most harmful to wolves, particularly pups, is Sarcoptes scabiei (or mange mite),[193] though they rarely develop full blown mange, unlike foxes.[11] Lice, such as Trichodectes canis, may cause sickness in wolves, but rarely death. Ticks of the genus Ixodes can infect wolves with Lyme disease and Rocky Mountain spotted fever.[193] The tick Dermacentor pictus also infests wolves. Other ectoparasites include biting lice, sucking lice and the fleas Pulex irritans and Ctenocephalides canis.[11]

Endoparasites known to infect wolves include protozoans and helminths (flukes, tapeworms, roundworms and thorny-headed worms). Of 30,000 protozoan species, only a few have been recorded to infect wolves: Isospora, Toxoplasma, Sarcocystis, Babesia, and Giardia.[193] Wolves may carry Neospora caninum, which is of particular concern to farmers, as the disease can be spread to livestock, with infected animals being 3–13 times more likely to miscarry than those not infected.[194][195] Among flukes, the most common in North American wolves is Alaria, which infects small rodents and amphibians, which are eaten by wolves. Upon reaching maturity, Alaria migrates to the wolf's intestine, but harms it little. Metorchis conjunctus, which enters wolves through eating fish, infects the wolf's liver or gall bladder, causing liver disease, inflammation of the pancreas, and emaciation. Most other fluke species reside in the wolf's intestine, though Paragonimus westermani lives in the lungs. Tapeworms are commonly found in wolves, as their primary hosts are ungulates, small mammals, and fish, which wolves feed upon. Tapeworms generally cause little harm in wolves, though this depends on the number and size of the parasites, and the sensitivity of the host. Symptoms often include constipation, toxic and allergic reactions, irritation of the intestinal mucosa, and malnutrition. Infections by the tapeworm Echinococcus granulosus in ungulate populations tend to increase in areas with high wolf densities, as wolves can shed Echinoccocus eggs in their feces onto grazing areas. Wolves can carry over 30 roundworm species, though most roundworm infections appear benign, depending on the number of worms and the age of the host. Ancylostoma caninum attaches itself on the intestinal wall to feed on the host's blood, and can cause hyperchromic anemia, emaciation, diarrhea, and possibly death. Toxocara canis, a hookworm known to infect wolf pups in utero, can cause intestinal irritation, bloating, vomiting, and diarrhea. Wolves may catch Dioctophyma renale from minks, which infects the kidneys, and can grow to lengths of 100 cm. D. renale causes the complete destruction of the kidney's functional tissue, and can be fatal if both kidneys are infected. Wolves can tolerate low levels of Dirofilaria immitis for many years without showing any ill effects, though high levels can kill wolves through cardiac enlargement and congestive hepatopathy. Wolves probably become infected with Trichinella spiralis by eating infected ungulates. Although T. spiralis isn't known to produce clinical signs in wolves, it can cause emaciation, salivation, and crippling muscle pains in dogs.[193] Thorny-headed worms rarely infect wolves, though three species have been identified in Russian wolves: Onicola skrjabini, Macrocantorhynchus catulinus, and Moniliformis moniliformis.[193]

Relationships with humans[edit]

Romulus and Remus nursed by the She-wolf (ca. 1616), Peter Paul Rubens

In culture[edit]

In mythology and folklore[edit]

Further information: Wulf, Big Bad Wolf and Lycanthropy

In Norse and Japanese mythology, wolves were portrayed as near deities: in Japan, grain farmers worshiped wolves at shrines and left food offerings near their dens, beseeching them to protect their crops from wild boars and deer,[196] while the wolf Fenrir of Norse mythology was depicted as the son of Loki. Other cultures portrayed wolves as part of their foundation myths: in the mythology of the Turks,[197] Mongols and Ainu, wolves were believed to be the ancestors of their people,[198] while the Dena’ina believed wolves were once men, and viewed them as brothers.[199] Wolves were linked to the sun in some Eurasian cultures: the Ancient Greeks and Romans associated wolves with the sun god Apollo,[199] while the wolf Sköll in Norse mythology was depicted pursuing the setting sun.[200] In Roman mythology, the Capitoline Wolf nurses Romulus and Remus, the future founders of Rome. According to the Pawnee creation myth, the wolf was the first animal to experience death.[201] Wolves were sometimes associated with witchcraft in both northern European and some Native American cultures: in Norse folkore, the völva (witch) Hyndla and the giantess Hyrrokin are both portrayed as using wolves as mounts, while in Navajo culture, wolves were feared as witches in wolf's clothing.[202] Similarly, the Tsilhqot'in believed that contact with wolves could cause mental illness and death.[199]

In fable and literature[edit]

One of the earliest written references to gray wolves occurs in the Babylonian epic Gilgamesh, in which the titular character rejects the sexual advances of the goddess Ishtar, reminding her that she had transformed a previous lover, a shepherd, into a wolf, thus turning him into the very animal that his flocks must be protected against.[203] According to the Avesta, the sacred text of the Zoroastrians, wolves are a creation of the evil spirit Ahriman, and are ranked among the most cruel of animals.[204] Aesop featured wolves in several of his fables, playing on the concerns of Ancient Greece's settled, sheep-herding world. His most famous is the fable of The Boy Who Cried Wolf, which is directed at those who knowingly raise false alarms, and from which the idiomatic phrase "to cry wolf" is derived. Some of his other fables concentrate on maintaining the trust between shepherds and guard dogs in their vigilance against wolves, as well as anxieties over the close relationship between wolves and dogs. Although Aesop used wolves to warn, criticize and moralize about human behaviour, his portrayals added to the wolf's image as a deceitful and dangerous animal.[205] This is mirrored in the Bible, where wolves are referenced thirteen times as symbols of greed and destructiveness.[206] Much of the symbolism Jesus used in the New Testament revolved around the pastoral culture of Israel, and explained his relationship with his followers as analogous to that of a good shepherd protecting his flock from wolves. An innovation in the popular image of wolves started by Jesus includes the concept of the wolf in sheep's clothing, which warns people against false prophets.[207] Several authors have proposed that Jesus's portrayal of wolves, comparing them to dangerous and treacherous people, was an important development in perceptions on the species, which legitimized centuries of subsequent wolf persecution in the western world.[207][208][209] Subsequent medieval Christian literature followed and expanded upon Biblical teachings on the wolf. It appeared in the seventh century edition of the Physiologus, which infused pagan tales with the spirit of Christian moral and mystical teaching. The Physiologus portrays wolves as being able to strike men dumb on sight, and of having only one cervical vertebra. Dante included a she-wolf, representing greed and fraud, in the first canto of the Inferno. The Malleus Maleficarum, first published in 1487, states that wolves are either agents of God sent to punish the wicked, or agents of Satan, sent with God's blessing to test the faith of believers.[210] Isengrim the wolf, a character first appearing in the 12th-century Latin poem Ysengrimus, is a major character in the Reynard Cycle, where he stands for the low nobility, whilst his adversary, Reynard the fox, represents the peasant hero. Although portrayed as loyal, honest and moral, Isengrim is forever the victim of Reynard's wit and cruelty, often dying at the end of each story.[211] The tale of Little Red Riding Hood, first written in 1697 by Charles Perrault, is largely considered to have had more influence than any other source of literature in forging the wolf's negative reputation in the western world. The wolf in this story is portrayed as a potential rapist, capable of imitating human speech.[212] The hunting of wolves, and their attacks on humans and livestock feature prominently in Russian literature, and are included in the works of Tolstoy, Chekhov, Nekrasov, Bunin, Sabaneyev, and others. Tolstoy's War and Peace and Chekhov's Peasants both feature scenes in which wolves are hunted with hounds and borzois.[213] Farley Mowat's largely fictional 1963 memoir Never Cry Wolf was the first positive portrayal of wolves in popular literature, and is largely considered to be the most popular book on wolves, having been adapted into a Hollywood film and taught in several schools decades after its publication. Although credited with having changed popular perceptions on wolves by portraying them as loving, cooperative and noble, it has been criticized for its idealization of wolves and its factual inaccuracies.[214][215][216]

In heraldry and symbolism[edit]

Main article: Wolves in heraldry
Emblem of the Serbian "White Wolves"

The wolf is a frequent charge in English armory. It is illustrated as a supporter on the shields of Lord Welby, Rendel, and Viscount Wolseley, and can be found on the coat of arms of Lovett and the vast majority of the Wilsons and Lows. The demi-wolf is a common crest, appearing in the arms and crests of members of many families, including that of the Wolfes, whose crest depicts a demi-wolf holding a crown in its paws, in reference to the assistance the family gave to Charles II during the battle of Worcester. Wolf heads are common in Scottish heraldry, particularly in the coats of Clan Robertson and Skene. The wolf is the most common animal in Spanish heraldry, and is often depicted as carrying a lamb in its mouth, or across its back.[217] The wolf is featured on the flags of the Confederated Tribes of the Colville Reservation, the Oneida Nation of Wisconsin and the Pawnee.[218] In modern times, the wolf is widely used as an emblem for military and paramilitary groups. It is the unofficial symbol of the spetsnaz, and serves as the logo of the Turkish Grey Wolves. During the Yugoslav Wars, several Serb paramilitary units adopted the wolf as their symbol, including the White Wolves and the Wolves of Vučjak.[219]

Livestock and dog predation[edit]

Waiting for a Chinook (ca. 1900), Charles Marion Russell

Livestock depredation has been one of the primary reasons for hunting wolves, and can pose a severe problem for wolf conservation: as well as causing economic losses, the threat of wolf predation causes great stress on livestock producers, and no foolproof solution of preventing such attacks short of exterminating wolves has been found.[220] Some nations help offset economic losses to wolves through compensation programmes or state insurance.[221] Domesticated animals are easy prey for wolves, as they have evolved under constant human protection, and are thus unable to defend themselves very well.[222] Wolves typically resort to attacking livestock when wild prey is depleted: in Eurasia, a large part of the diet of some wolf populations consists of livestock, while such incidences are rare in North America, where healthy populations of wild prey have been largely restored.[220] The majority of losses occur during the summer grazing period, with untended livestock in remote pastures being the most vulnerable to wolf predation.[223] The most frequently targeted livestock species are sheep (Europe), domestic reindeer (northern Scandinavia), goats (India), horses (Mongolia), cattle and turkeys (North America).[220] The number of animals killed in single attacks varies according to species: most attacks on cattle and horses result in one death, while turkeys, sheep and domestic reindeer may be killed in surplus.[224] Wolves mainly attack livestock when the animals are grazing, though they occasionally break into fenced enclosures.[112] In some cases, wolves do not need to physically attack livestock to negatively affect it: the stress livestock experiences in being vigilant for wolves may result in miscarriages, weight loss and a decrease in meat quality.[194]

Wolves kill dogs on occasion, with some wolf populations relying on dogs as an important food source.[220] In Croatia, wolves kill more dogs than sheep, and wolves in Russia appear to limit stray dog populations. Wolves may display unusually bold behaviour when attacking dogs accompanied by people, sometimes ignoring nearby humans.[220] Wolf attacks on dogs may occur both in house yards and in forests.[225] Wolf attacks on hunting dogs are considered a major problem in Scandinavia and Wisconsin.[220][225] The most frequently killed hunting breeds in Scandinavia are harriers, with older animals being most at risk, likely because they are less timid than younger animals, and react to the presence of wolves differently. Large hunting dogs such as Swedish elkhounds are more likely to survive wolf attacks due to their better ability to defend themselves.[225]

Attacks on humans[edit]

Child snatched by a wolf (1914), Le Petit Journal

The fear of wolves has been pervasive in many societies, though humans are not part of the wolf's natural prey.[24] How wolves react to humans depends largely on their prior experience with people: wolves lacking any negative experience of humans, or are food conditioned, may show little fear of people.[226] Although wolves may react aggressively under provocation, such attacks are mostly limited to quick bites on extremities, and the attacks are not pressed. Predatory attacks (attacks by wolves treating humans as food) may be preceded by a long period of habituation, in which wolves gradually lose their fear of humans. The victims are repeatedly bitten on the head and face, and are then dragged off and consumed, unless the wolves are driven off. Such attacks typically occur only locally, and do not stop until the wolves involved are eliminated. Predatory attacks can occur at any time of the year, with a peak in the June–August period, when the chances of people entering forested areas (for livestock grazing or berry and mushroom picking) increase,[24] though cases of non-rabid wolf attacks in winter have been recorded in Belarus, the Kirovsk and Irkutsk districts, Karelia and Ukraine.[11] Also, wolves with pups experience greater food stresses during this period. The majority of victims of predatory wolf attacks are children under the age of 18 and, in the rare cases where adults are killed, the victims are almost always women. Cases of rabid wolves are low when compared to other species, as wolves do not serve as primary reservoirs of the disease, but can be infected by animals such as dogs, jackals and foxes. Incidents of rabies in wolves are very rare in North America, though numerous in the eastern Mediterranean, Middle East and Central Asia. Wolves apparently develop the "furious" phase of rabies to a very high degree which, coupled with their size and strength, makes rabid wolves perhaps the most dangerous of rabid animals,[24] with bites from rabid wolves being 15 times more dangerous than those of rabid dogs.[11] Rabid wolves usually act alone, travelling large distances and often biting large numbers of people and domestic animals. Most rabid wolf attacks occur in the spring and autumn periods. Unlike with predatory attacks, the victims of rabid wolves are not eaten, and the attacks generally only occur on a single day. Also, the victims are chosen at random, though the majority of cases involve adult men. In the half-century up to 2002, there were eight fatal attacks in Europe and Russia, and more than 200 in south Asia.[24] Between 2005-2010, two people were killed in North America.[227][228]

Hunting[edit]

Carcasses of hunted wolves in Kamyshinsky District, Volgograd Oblast, Russia.

Wolves are notoriously difficult to hunt due to their elusiveness, their sharp senses, their high endurance in the chase and ability to quickly incapacitate and kill hunting dogs.[229] Historically, many methods have been devised to hunt wolves, including the killing of spring-born litters in their dens, coursing with dogs (usually combinations of sighthounds, bloodhounds and fox terriers), poisoning with strychnine, and foothold and deadfall traps.[230][231][232] A popular method of wolf hunting in Russia involves trapping a pack within a small area by encircling it with fladry poles carrying a human scent. This method relies heavily on the wolf's fear of human scents, though it can lose its effectiveness when wolves become accustomed to the smell.[232] Some hunters are able to lure wolves by imitating their calls.[232] In Kazakhstan and Mongolia, wolves are traditionally hunted with eagles and falcons, though this practise is declining, as experienced falconers are becoming few in number.[232] Shooting wolves from aircraft is highly effective, as it allows greater visibility of wolves than hunting on the ground,[232] though this method is controversial, as it allows wolves little chance to escape or defend themselves.[233]

Uses[edit]

Gray wolf skins at the Bergen fishmarket, Norway.

While not in the same class as high grade furbearers like beaver, otter or mink, the gray wolf's fur is nonetheless thick and durable,[234] and is primarily used for scarfs and the trimmings of women's garments, though its occasionally used for jackets, short capes, coats,[235] mukluks and rugs.[57] Aside from bodily protection and adornment, gray wolf pelts have also been used as camouflage in hunting and warfare, as an insignia among western Native Americans and as a form of currency.[234] The quality of wolf peltries rests on the density and strength of the fur fibre, which keeps the fur upright and gives the pelt an appealing bushy aspect. These characteristics are mostly found in northern wolf populations, but gradually lessen further south in warmer climates. North American wolf pelts are among the most valuable, as they are silkier and fluffier than Eurasian peltries.[235] In Medieval Europe, pelts were considered the only practical aspect of wolves, though they were seldom used, due to the skin's foul odour.[236] In Scandinavian folklore, wolf-skin girdles assisted in transforming the wearers into werewolves,[237] while several Native American tribes used wolf pelts for medicinal purposes.[231] Plains Indians often wore wolf pelts as disguises to get close to American bison when hunting.[238] The Pawnee wore wolf skins as capes when exploring enemy territories.[239] The United States Army used wolf skin for parkas during the later stages of World War II and the Korean War to protect the faces of soldiers from frostbite.[235] In the Soviet Union, 30,000 wolf pelts were produced annually between 1976 and 1988. Statistics from CITES indicate that 6,000–7,000 wolf skins are internationally traded each year, with Canada, the former Soviet Union, Mongolia and China being the largest exporters, and the United States and Great Britain being the largest importers. Overall, the harvesting of wolves for their fur has little impact on their population, as only the northern varieties (whose numbers are stable) are of commercial value.[240] Wolf trapping for fur remains a lucrative source of income for many Native Americans.[57]

It is rare for wolves to be hunted for food, though historically, people have resorted to consuming wolf flesh in times of scarcity, or for medicinal reasons. Wolf meat was eaten several times during Vilhjalmur Stefansson's 1913 Arctic expedition, particularly during the summer, when wolves were fat.[241] Natives in Transbaikalia reportedly ate wolf meat even when food was plentiful.[242] Most Native American tribes, particularly the Naskapis, viewed wolf flesh as edible but inadequate nutrition, as it was not a herbivore and thus did not possess the same healing qualities thought to be distinct in plant eaters.[239] The consumption of wolf flesh and organs plays an important role in Asian folk medicine: in Mongolia, eating the meat and lungs of a wolf is said to alleviate colds, and sprinkling food with powdered wolf rectum is said to cure haemorroids.[243] Some Japanese mountain people ate wolf meat to give them courage.[244] During the filming of The Grey, the cast members famously ate wolf meat.[245] Accounts on how wolf meat tastes vary greatly, with descriptions ranging from "tough",[244] "gristly",[246] "distasteful" and "smelly",[242] to "somewhat [resembling] chicken",[247] and "very superior to lean venison".[248]

Gray wolves as pets and working animals[edit]

Gray wolves and coyotes used as draught animals in northern Ontario, 1923.

American biologist, Stanley P. Young, described tame wolves as thus:

Generally speaking, on the basis of their experience, tame wolves are strictly "one-man dogs". They may be confiding and playful with the man who raised them, or even with his whole family, if fed and cared for by them, but they are suspicious and timid in the presence of strangers. They invariably retain certain reactions of wolf nature, as for instance, an incorrigible desire to kill chickens or other small livestock whenever opportunity arises.[249]

Several hunters of the USFWS kept wolf pups as pets, with the best results occurring when they were caught just after their eyes began opening. In contrast, pups taken at 3–4 weeks of age proved unmanageable, with only one in 11 of such pups becoming tame, despite one month of eight hours per day of socialization with people.[249] Though wolves are trainable, they lack the same degree of tractability seen in dogs. Most attempts to train wolves as working dogs have met with failure. Swedish biologist Erik Zimen attempted to train his captive wolves as sled dogs, and although his wolves eventually accepted the harness and the need to pull the sled in a straight line, they were ultimately unreliable, as they fought for personal space, would ignore commands once tired, and were distracted by other wildlife.[250] However, John James Audubon recorded an instance of a wolf being trained to hunt deer in Kentucky,[251] and Henry Wharton Shoemaker published a similar account of settlers in western and central Pennsylvania using wolves as hunting dogs.[252] Buffon wrote in his Natural History of tamed wolves in Persia being trained to perform dances and tricks.[253]

See also[edit]

Notes[edit]

  1. ^ The English 'wolf' stems from the Old English wulf, which is itself thought to be derived from the Proto-Germanic *wulfaz, from the Proto-Indo-European root *wlqwos/*lukwos.[3] Old English literature contains several instances of Anglo-Saxon kings and warriors taking on wulf as a prefix or suffix in their names. Examples include Wulfhere, Cynewulf, Ceonwulf, Wulfheard, Earnwulf, Wulfmǣr, Wulfstān and Æthelwulf. Wolf-related names were also common among pre-Christian Germanic warriors: Wolfhroc (Wolf-Frock), Wolfhetan (Wolf Hide), Isangrim (Grey Mask), Scrutolf (Garb Wolf), Wolfgang (Wolf Gait) and Wolfdregil (Wolf Runner).[4]
  2. ^ The species also includes the domestic dog (Canis lupus familiaris) and the dingo (Canis lupus dingo), both of which are man-made variants. However, neither the dog nor the dingo would normally be referred to as wolves. Throughout this article, the term "gray wolf" will often be used to refer collectively to naturally ocurring subspecies, especially the nominate subspecies, Canis lupus lupus.
  3. ^ The term "western wolf" is primarily used by taxonomists in distinguishing Canis lupus from Canis lycaon, the so-called "eastern wolf".[8]
  4. ^ In the past, the prevailing view on gray wolf packs was that they consisted of individuals vying with each other for dominance, with dominant gray wolves being referred to as the "alpha" male and female, and the subordinates as "beta" and "omega" wolves. This terminology was first used in 1947 by Rudolf Schenkel of the University of Basel, who based his findings on researching the behavior of captive gray wolves. This view on gray wolf pack dynamics was later popularized by L. David Mech in his 1970 book The Wolf. He formally disavowed this terminology in 1999, explaining that it was heavily based on the behavior of captive packs consisting of unrelated individuals, an error reflecting the once prevailing view that wild pack formation occurred in winter among independent gray wolves. Later research on wild gray wolves revealed that the pack is usually a family consisting of a breeding pair and its offspring of the previous 1–3 years.[81]

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