Moose

Etymology

Alces alces is called a "moose" in North American English, but an "elk" in British English.^[3] The word "elk" in North American English refers to a completely different species of deer, Cervus canadensis, also called the wapiti (from Algonquin). A mature male moose is called a bull, a mature female a cow, and an immature moose of either sex a calf.

In Classical Antiquity, the animal was known as ἄλκη álkē^[4] in Greek and alces^[5] in Latin, words probably borrowed from a Germanic language or another language of northern Europe.^[3] By the 8th century, during the Early Middle Ages, the species was known as Old English: elch, elh, eolh, derived from the Proto-Germanic: *elho-, *elhon- and possibly connected with the Old Norse: elgr.^[3] Later, the species became known in Middle English as elk, elcke, or elke, appearing in the Latinized form alke, with the spelling alce borrowed directly from Latin: alces.^[3][6]

The word "elk" remained in usage because of English-speakers' familiarity with the species in Continental Europe; however, without any living animals around to serve as a reference, the meaning became rather vague, and by the 17th century "elk" had a meaning similar to "large deer".^[7] Dictionaries of the 18th century simply described "elk" as a deer that was "as large as a horse".^[8]

The word "moose" had first entered English by 1606^[9] and is borrowed from the Algonquian languages (compare the Narragansett moos and Eastern Abenaki mos; according to early sources, these were likely derived from moosu, meaning "he strips off"),^[10] and possibly involved forms from multiple languages mutually reinforcing one another. The Proto-Algonquian form was *mo·swa.^[11]

Antlers

Bull moose have antlers like other members of the deer family. The size and growth rate of antlers is determined by diet and age.^[20] Size and symmetry in the number of antler points signals bull moose health and cows may select mates based on antler size and symmetry.^[21] Bull moose use their antlers to display dominance, to discourage competition, and to spar or fight rivals.^[20]

The male's antlers grow as cylindrical beams projecting on each side of the head at right angles to the midline of the skull, and then fork. The lower prong of this fork may be either simple, or divided into two or three tines, with some flattening. Most moose have antlers that are broad and palmate (flat) with tines (points) along the outer edge.^[20] Within the ecologic range of the moose in Europe, northern populations display the palmate pattern of antlers, while the antlers of European moose residing the southerly portion of its range are typically of the cervina dendritic pattern and comparatively small, perhaps due to evolutionary pressures of hunting by humans, who prize the large palmate antlers. European moose with antlers intermediate between the palmate and the dendritic form are found in the middle of the north-south range.^[22] Moose with antlers have more acute hearing than those without antlers; a study of trophy antlers using a microphone found that the palmate antler acts as a parabolic reflector, amplifying sound at the moose's ear.^[23]

The antlers of mature Alaskan adult bull moose (5 to 12 years old) have a normal maximum spread greater than 200 centimeters (79 in). By the age of 13, moose antlers decline in size and symmetry. The widest spread recorded was 210 centimeters (83 in) across. An Alaskan moose also holds the record for the heaviest weight at 36 kilograms (79 lb).^[20]

Antler beam diameter, not the number of tines, indicates age.^[20] In North America, moose (A. a. americanus) antlers are usually larger than those of Eurasian moose and have two lobes on each side, like a butterfly. Eurasian moose antlers resemble a seashell, with a single lobe on each side.^[20] In the North Siberian moose (A. a. bedfordiae), the posterior division of the main fork divides into three tines, with no distinct flattening. In the common moose (A. a. alces) this branch usually expands into a broad palmation, with one large tine at the base and a number of smaller snags on the free border. There is, however, a Scandinavian breed of the common moose in which the antlers are simpler and recall those of the East Siberian animals. The palmation appears to be more marked in North American moose than in the typical Scandinavian moose.

After the mating season males drop their antlers to conserve energy for the winter. A new set of antlers will then regrow in the spring. Antlers take three to five months to fully develop, making them one of the fastest growing animal organs. Antler growth is "nourished by an extensive system of blood vessels in the skin covering, which contains numerous hair follicles that give it a 'velvet' texture."^[20] This requires intense grazing on a highly-nutritious diet. By September the velvet is removed by rubbing and thrashing which changes the colour of the antlers. Immature bulls may not shed their antlers for the winter, but retain them until the following spring. Birds, carnivores and rodents eat dropped antlers as they are full of protein and moose themselves will eat antler velvet for the nutrients.^[20]

If a bull moose is castrated, either by accidental or chemical means, he will shed his current set of antlers within two weeks and then immediately begin to grow a new set of misshapen and deformed antlers that he will wear the rest of his life without ever shedding again; similarly deformed antlers can result from a deficiency of testosterone caused by cryptorchidism or old age.^[24] These deformed antlers are composed of living bone which is still growing or able to grow, since testosterone is needed to stop antler growth;^[25] they may take one of two forms. "Cactus antlers" or velericorn antlers^[25] usually retain the approximate shape of a normal moose's antlers but have numerous pearl-shaped exostoses on their surface; being made of living bone, they are easily broken but can grow back. Perukes (US: /pəˈruːks/) are constantly growing, tumor-like antlers with a distinctive appearance similar to coral.^[24] Like roe deer, moose are more likely to develop perukes, rather than cactus antlers, than the more developed cervine deer, but unlike roe deer, moose do not suffer fatal decalcification of the skull as a result of peruke growth,^[26][25] but rather can support their continued growth until they become too large to be fully supplied with blood.^[24] The distinctive-looking perukes (often referred to as "devil's antlers") are the source of several myths and legends among many groups of Inuit as well as several other tribes of indigenous peoples of North America.

In extremely rare circumstances, a cow moose may grow antlers. This is usually attributed to a hormone imbalance.^[27]

Proboscis and olfaction

The moose proboscis is distinctive among living cervids due to its large size; it also features nares that can be sealed shut when the moose is browsing aquatic vegetation. The moose proboscis likely evolved as an adaptation to aquatic browsing, with loss of the rhinarium, and development of a superior olfactory column separate from an inferior respiratory column.^[28] This separation contributes to the moose's keen sense of smell, which they employ to detect water sources, to find food under snow, and to detect mates or predators.^[29][28]

Hooves

As with all members of the order Artiodactyla (even-toed ungulates), moose feet have two large keratinized hooves corresponding to the third and fourth toe, with two small posterolateral dewclaws (vestigial digits), corresponding to the second and fifth toe. The hoof of the fourth digit is broader than that of the third digit, while the inner hoof of the third digit is longer than that of the fourth digit. This foot configuration may favor striding on soft ground.^[30] The moose hoof splays under load, increasing surface area, which limits sinking of the moose foot into soft ground or snow, and which increases efficiency when swimming. The body weight per footprint surface area of the moose foot is intermediate between that of the pronghorn foot, (which have stiff feet lacking dewclaws—optimized for high-speed running) and the caribou foot (which are more rounded with large dewclaws, optimized for walking in deep snow). The moose's body weight per surface area of footprint is about twice that of the caribou.^[31][32]

Skin and fur

Moose skin is typical of the deer family. Moose fur consists of four types of hair: eyelashes, whiskers, guard hairs and wool hairs. Hair length and hair density varies according to season, age, and body region.^[33] The coat has two layers—a top layer of long guard hairs and a soft wooly undercoat. The guard hairs are hollow and filled with air for better insulation, which also helps them stay afloat when swimming.^[34][35]

Dewlap

Both male and female moose have a dewlap or bell,^[36] which is a fold of skin under the chin. Its exact function is unknown, but some morphologic analyses suggest a cooling (thermoregulatory) function.^[37] Other theories include a fitness signal in mating, as a visual and olfactory signal, or as a dominance signal by males, as are the antlers.^[38]

Diet

The moose is a browsing herbivore and is capable of consuming many types of plant or fruit. The average adult moose needs to consume 96 megajoules (23,000 kilocalories) per day to maintain its body weight.^[39] Much of a moose's energy is derived from terrestrial vegetation, mainly consisting of forbs and other non-grasses, and fresh shoots from trees such as willow and birch. As these terrestrial plants are rather low in sodium, as much as half of its diet usually consists of aquatic plants, including lilies and pondweed,^[40] which while lower in energy content, provide the moose with its sodium requirements.^[41] In winter, moose are often drawn to roadways, to lick salt that is used as a snow and ice melter.^[42] A typical moose, weighing 360 kg (794 lb), can eat up to 32 kg (71 lb) of food per day.^[41]

Moose lack upper front teeth, but have eight sharp incisors on the lower jaw. They also have a tough tongue, lips and gums, which aid in the eating of woody vegetation. Moose have six pairs of large, flat molars and, ahead of those, six pairs of premolars, to grind up their food. A moose's upper lip is very sensitive, to help distinguish between fresh shoots and harder twigs, and is prehensile, for grasping their food. In the summer, moose may use this prehensile lip for grabbing branches and pulling, stripping the entire branch of leaves in a single mouthful, or for pulling forbs, like dandelions, or aquatic plants up by the base, roots and all.^[43][44] A moose's diet often depends on its location, but they seem to prefer the new growths from deciduous trees with a high sugar content, such as white birch, trembling aspen and striped maple, among many others.^[45] To reach high branches, a moose may bend small saplings down, using its prehensile lip, mouth or body. For larger trees a moose may stand erect and walk upright on its hind legs, allowing it to reach branches up to 4.26 meters (14 ft 0 in) or higher above the ground.^[46][47]

Moose are excellent swimmers and are known to wade into water to eat aquatic plants. This trait serves a second purpose in cooling down the moose on summer days and ridding itself of black flies. Moose are thus attracted to marshes and river banks during warmer months as both provide suitable vegetation to eat and water to wet themselves in. Moose have been known to dive over 5.5 metres (18 ft) to reach plants on lake bottoms,^[48] and the complex snout may assist the moose in this type of feeding. Moose are the only deer that are capable of feeding underwater.^[49] As an adaptation for feeding on plants underwater, the nose is equipped with fatty pads and muscles that close the nostrils when exposed to water pressure, preventing water from entering the nose.^[50] Other species can pluck plants from the water too, but these need to raise their heads in order to swallow.

This fenced-in area is part of a long-term research project to examine the effects of moose browsing on plant biodiversity.

Moose are not grazing animals but browsers (concentrate selectors). Like giraffes, moose carefully select foods with less fiber and more concentrations of nutrients. Thus, the moose's digestive system has evolved to accommodate this relatively low-fiber diet. Unlike most hooved, domesticated animals (ruminants), moose cannot digest hay, and feeding it to a moose can be fatal.^[51][52] The moose's varied and complex diet is typically expensive for humans to provide, and free-range moose require a lot of forested hectarage for sustainable survival, which is one of the main reasons moose have never been widely domesticated.^{[citation needed]}

• 
  Bark stripping
• 
  Bull moose eating a fireweed plant
• 
  Bull moose browses a beaver pond

Natural predators

A full-grown moose has few enemies except Siberian tigers (Panthera tigris tigris) which regularly prey on adult moose,^[53][54][55] but a pack of gray wolves (Canis lupus) can still pose a threat, especially to females with calves.^[56] Brown bears (Ursus arctos)^[17] are also known to prey on moose of various sizes and are the only predator besides the wolf to attack moose both in Eurasia and North America. In Western Russia, moose provide about 15% annual estimated dietary energy content for brown bears and are the most important food source for these predators during spring.^[57] However, Brown bears are more likely to scavenge a wolf kill or to take young moose than to hunt adult moose on their own.^[58][59][60] Black bears (Ursus americanus) and cougars (Puma concolor) can be significant predators of moose calves in May and June and can, in rare instances, prey on adults (mainly cows rather than the larger bulls).^[61][62] Wolverines (Gulo gulo) are most likely to eat moose as carrion but have killed moose, including adults, when the large ungulates are weakened by harsh winter conditions.^[63][64][65] Orcas (Orcinus orca) are the moose's only confirmed marine predator as they have been known to prey on moose and other deer swimming between islands out of North America's Northwest Coast. However, such kills are rare and a matter of opportunity, as moose are not a regular part of the orca diet.^[66] There is at least one recorded instance of a moose being scavenged by a Greenland shark (Somniosus microcephalus).^[67]

In some areas, moose are the primary source of food for wolves. Moose usually flee upon detecting wolves. Wolves usually follow moose at a distance of 100 to 400 meters (300 to 1,300 ft), occasionally at a distance of two to three kilometers (1 to 2 mi). Attacks from wolves against young moose may last seconds, though sometimes they can be drawn out for days with adults. Sometimes, wolves will chase moose into shallow streams or onto frozen rivers, where their mobility is greatly impeded. Moose will sometimes stand their ground and defend themselves by charging at the wolves or lashing out at them with their powerful hooves. Wolves typically kill moose by tearing at their haunches and perineum, causing massive blood loss. Occasionally, a wolf may immobilize a moose by biting its sensitive nose, the pain of which can paralyze a moose.^[68] Wolf packs primarily target calves and elderly animals, but can and will take healthy, adult moose. Moose between the ages of two and eight are seldom killed by wolves.^[69] Though moose are usually hunted by packs, there are cases in which single wolves have successfully killed healthy, fully-grown moose.^[70][71]

Research into moose predation suggests that their response to perceived threats is learned rather than instinctual. In practical terms this means moose are more vulnerable in areas where wolf or bear populations were decimated in the past but are now rebounding. These same studies suggest, however, that moose learn quickly and adapt, fleeing an area if they hear or smell wolves, bears, or scavenger birds such as ravens.^[72]

Moose are also subject to various diseases and forms of parasitism. In northern Europe, the moose botfly is a parasite whose range seems to be spreading.^[73]

Parasites

Moose typically carry a heavy burden of parasites, both externally and internally. Parasitosis is an important cause of moose morbidity and mortality and also contributes to vulnerability to predators.^[74] Ectoparasites of moose include the moose nose bot fly,^[75] and winter ticks.^[76] Endoparasites of moose include dog tapeworm, meningeal worm,^[77] lungworm, and roundworm.

Social structure and reproduction

Moose are mostly diurnal. They are generally solitary with the strongest bonds between mother and calf. Although moose rarely gather in groups, there may be several in close proximity during the mating season.

Rutting and mating occurs in September and October. During the rut, mature bulls will cease feeding completely for a period of approximately two weeks; this fasting behavior has been attributed to neurophysiological changes related to redeployment of olfaction for detection of moose urine and moose cows.^[78] The males are polygynous and will seek several females to breed with. During this time both sexes will call to each other. Males produce heavy grunting sounds that can be heard from up to 500 metres (1,600 ft) away, while females produce wail-like sounds.^[79] Males will fight for access to females. Initially, the males assess which of them is dominant and one bull may retreat, however, the interaction can escalate to a fight using their antlers.

Female moose have an eight-month gestation period, usually bearing one calf, or twins if food is plentiful,^[80] in May or June.^[81] Twinning can run as high as 30% to 40% with good nutrition^[82] Newborn moose have fur with a reddish hue in contrast to the brown appearance of an adult. The young will stay with the mother until just before the next young are born. The life span of an average moose is about 15–25 years. Moose populations are stable at 25 calves for every 100 cows at 1 year of age. With availability of adequate nutrition, mild weather, and low predation, moose have a huge potential for population expansion.^[82]

• 
  (newborn)
  Calves nursing in spring.
• 
  (3 months)
  Calves stay near their mothers at all times.
• 
  (9 months)
  This calf is almost ready to leave its mother.
• 
  (10–11 months)
  This yearling was probably recently chased away by its pregnant mother.

Aggression

Moose are not typically aggressive towards humans, but will be aggressive when provoked or frightened. Moose attack more people than bears and wolves combined, but usually with only minor consequences. In the Americas, moose injure more people than any other wild mammal; worldwide, only hippopotamuses injure more.^[83] When harassed or startled by people or in the presence of a dog, moose may charge. Also, as with bears or most wild animals, moose accustomed to being fed by people may act aggressively when denied food. During the fall mating season, bulls may be aggressive toward humans. Cows are protective of young calves and will attack humans who come close, especially if they come between mother and calf. Moose are not territorial, do not view humans as food, and usually will not pursue humans who run away.^[84]

Moose are unpredictable. They are most likely to attack if annoyed or harassed, or if approached too closely. A moose that has been harassed may vent its anger on anyone in the vicinity, and they often do not make distinctions between their tormentors and innocent passersby.^{[citation needed]} Moose are very limber animals with highly flexible joints and sharp, pointed hooves, and are capable of kicking with both front and back legs. Unlike other large, hoofed mammals, such as horses, moose can kick in all directions, including sideways. Thus, there is no safe side from which to approach. Moose often give warning signs prior to attacking, displaying aggression by means of body language. Maintained eye contact is usually the first sign of aggression, while laid-back ears or a lowered head is a sign of agitation. When the hairs on the back of the moose's neck and shoulders (hackles) stand up, a charge is usually imminent. The Anchorage Visitor Centers warn tourists that "...a moose with its hackles raised is a thing to fear."^{[85][86][87][88]}

Moose cows are more likely to emit protest moans when courted by small males. This attracts the attention of large males, promotes male-male competition and violence, reduces harassment of cows by small males, and increases mating opportunities with large males.^[89] This in turn means that the cow moose has at least a small degree of control over which bulls she mates with.^[90]

Moose often show aggression to other animals as well, especially predators. Bears are common predators of moose calves and, rarely, adults. Alaskan moose have been reported to successfully fend off attacks from both black and brown bears. Moose have been known to stomp attacking wolves, which makes them less preferred as prey to the wolves. Moose are fully capable of killing bears and wolves. In one rare event, a female moose killed two adult male wolves.^[91][92] A moose of either sex that is confronted by danger may let out a loud roar, more resembling that of a predator than a prey animal. European moose are often more aggressive than North American moose, such as the moose in Sweden, which often become very agitated at the sight of a predator. However, like all ungulates known to attack predators, the more aggressive individuals are always darker in color, with the darkest coloring usually in areas facing the opponent, thus serving as a natural warning to other animals.^[49]

Habitat

Moose require habitat with adequate edible plants (e.g., pond grasses, young trees and shrubs), cover from predators, and protection from extremely hot or cold weather. Moose travel among different habitats with the seasons to address these requirements.^[93] Moose are cold-adapted mammals with thickened skin, dense, heat-retaining coat, and a low surface:volume ratio, which provides excellent cold tolerance but poor heat tolerance. Moose survive hot weather by accessing shade or cooling wind, or by immersion in cool water. In hot weather, moose are often found wading or swimming in lakes or ponds. When heat-stressed, moose may fail to adequately forage in summer and may not gain adequate body fat to survive the winter. Also, moose cows may not calve without adequate summer weight gain. Moose require access to both young forest for browsing and mature forest for shelter and cover. Forest disturbed by fire and logging promotes the growth of fodder for moose. Moose also require access to mineral licks, safe places for calving and aquatic feeding sites.^[93]

Moose avoid areas with little or no snow as this increases the risk of predation by wolves and avoid areas with deep snow, as this impairs mobility. Thus, moose select habitat on the basis of trade-offs between risk of predation, food availability, and snow depth.^[94] With reintroduction of bison into boreal forest, there was some concern that bison would compete with moose for winter habitat, and thereby worsen the population decline of moose. However, this does not appear to be a problem. Moose prefer sub-alpine shrublands in early winter, while bison prefer wet sedge valley meadowlands in early winter. In late winter, moose prefer river valleys with deciduous forest cover or alpine terrain above the tree line, while bison preferred wet sedge meadowlands or sunny southern grassy slopes.^[95]

North America

After expanding for most of the 20th century, the moose population of North America has been in steep decline since the 1990s. Populations expanded greatly with improved habitat and protection, but now the moose population is declining rapidly.^[96] This decline has been attributed to opening of roads and landscapes into the northern range of moose, allowing deer to become populous in areas where they were not previously common. This encroachment by deer on moose habitat brought moose into contact with previously unfamiliar pathogens, including brainworm and liver fluke, and these parasites are believed to have contributed to the population decline of moose.^[97][98]

In North America, the moose range includes almost all of Canada (excluding the arctic and Vancouver Island), most of Alaska, northern and eastern North Dakota,^[99] northern New England, the Adirondack Mountain region and Taconic highlands of northeast New York State,^[100][101] the upper Rocky Mountains, northern Minnesota, northern Wisconsin, Michigan's Upper Peninsula, and Isle Royale in Lake Superior. This massive range, containing diverse habitats, contains four of the six North American subspecies. In the West, moose populations extend across Canada (British Columbia and Alberta). Isolated groups have been verified as far south as the mountains of Utah and Colorado and as far west as the Lake Wenatchee area of the Washington Cascades.^[102][103] In the northwestern US, the range includes Wyoming, Montana, Idaho, and smaller areas of Washington, and Oregon.^[104] Moose have extended their range southwards in the western Rocky Mountains, with initial sightings in Yellowstone National Park in 1868, and then to the northern slope of the Uinta Mountains in Utah in the first half of the twentieth century.^[105] This is the southernmost naturally established moose population in the United States.^[105] In 1978, a few breeding pairs were reintroduced in western Colorado, and the state's moose population is now more than 2,400.^[106]

In northeastern North America, the Eastern moose's history is very well documented: moose meat was a staple in the diet of indigenous peoples for centuries. The common name "moose" was brought into English from the word used by those who lived in present day coastal Rhode Island. The indigenous people often used moose hides for leather and its meat as an ingredient in pemmican, a type of dried jerky used as a source of sustenance in winter or on long journeys.^[107]

The historical range of the subspecies extended from well into Quebec, the Maritimes, and Eastern Ontario south to include all of New England finally ending in the very northeastern tip of Pennsylvania in the west, cutting off somewhere near the mouth of the Hudson River in the south. The moose has been extinct in much of the eastern U.S. for as long as 150 years, due to colonial era overhunting and destruction of its habitat: Dutch, French, and British colonial sources all attest to its presence in the mid 17th century from Maine south to areas within 160 kilometers (100 mi) of present-day Manhattan. However, by the 1870s, only a handful of moose existed in this entire region in very remote pockets of forest; less than 20% of suitable habitat remained.^[108]

Since the 1980s, however, moose populations have rebounded, thanks to regrowth of plentiful food sources,^[108] abandonment of farmland, better land management, clean-up of pollution, and natural dispersal from the Canadian Maritimes and Quebec. South of the Canada–US border, Maine has most of the population with a 2012 headcount of about 76,000 moose.^[109] Dispersals from Maine over the years have resulted in healthy, growing populations each in Vermont and New Hampshire, notably near bodies of water and as high up as 910 m (3,000 ft) above sea level in the mountains. In Massachusetts, moose had gone extinct by 1870, but re-colonized the state in the 1960s, with the population expanding from Vermont and New Hampshire; by 2010, the population was estimated at 850–950.^[110] Moose reestablished populations in eastern New York and Connecticut and appeared headed south towards the Catskill Mountains, a former habitat.^{[111][112][113][114][115][116][117]}

In the Midwest U.S., moose are primarily limited to the upper Great Lakes region, but strays, primarily immature males, have been found as far south as eastern Iowa.^[118] For unknown reasons, the moose population is declining rapidly in the Midwest.^[96]

Moose were successfully introduced on Newfoundland in 1878 and 1904,^[119] where they are now the dominant ungulate, and somewhat less successfully on Anticosti Island in the Gulf of Saint Lawrence.^{[citation needed]}

• 
  Cow and bull moose
• 
  Cow moose
• 
  Moose calf

Europe and Asia

In Europe, moose are currently found in large numbers throughout Norway, Sweden, Finland, Latvia, Estonia, Poland, with more modest numbers in the southern Czech Republic, Belarus, and northern Ukraine. They are also widespread through Russia on up through the borders with Finland south towards the border with Estonia, Belarus and Ukraine and stretching far away eastwards to the Yenisei River in Siberia. The European moose was native to most temperate areas with suitable habitat on the continent and even Scotland from the end of the last Ice Age, as Europe had a mix of temperate boreal and deciduous forest. Up through Classical times, the species was certainly thriving in both Gaul and Magna Germania, as it appears in military and hunting accounts of the age. However, as the Roman era faded into medieval times, the beast slowly disappeared: soon after the reign of Charlemagne, the moose disappeared from France, where its range extended from Normandy in the north to the Pyrenees in the south. Farther east, it survived in Alsace and the Netherlands until the 9th century as the marshlands in the latter were drained and the forests were cleared away for feudal lands in the former. It was gone from Switzerland by the year 1000, from the western Czech Republic by 1300, from Mecklenburg in Germany by c. 1600, and from Hungary and the Caucasus since the 18th and 19th century, respectively.

By the early 20th century, the last strongholds of the European moose appeared to be in Fennoscandian areas and patchy tracts of Russia, with a few migrants found in what is now Estonia and Lithuania. The USSR and Poland managed to restore portions of the range within its borders (such as the 1951 reintroduction into Kampinos National Park and the later 1958 reintroduction in Belarus), but political complications limited the ability to reintroduce it to other portions of its range. Attempts in 1930 and again in 1967 in marshland north of Berlin were unsuccessful. At present in Poland, populations are recorded in the Biebrza river valley, Kampinos, and in Białowieża Forest. It has migrated into other parts of Eastern Europe and has been spotted in eastern and southern Germany.^[133][134] Unsuccessful thus far in recolonizing these areas via natural dispersal from source populations in Poland, Belarus, Ukraine, Czech Republic, and Slovakia, it appears to be having more success migrating south into the Caucasus. It is listed under Appendix III of the Bern Convention.^[135][136]

In 2008, two moose were reintroduced into the Scottish Highlands^[137][138] in Alladale Wilderness Reserve.^[139] The moose disappeared as a breeding species from Denmark about 4,500 years ago (in the last century, a very small number have lived for periods in Zealand without establishing a population after swimming across the Øresund from Sweden),^[140] but in 2016-17 ten were introduced to Lille Vildmose from Sweden. In 2020, this population had increased to about 25 animals.^[141]

The East Asian moose populations confine themselves mostly to the territory of Russia, with much smaller populations in Mongolia and Northeastern China. Moose populations are relatively stable in Siberia and increasing on the Kamchatka Peninsula. In Mongolia and China, where poaching took a great toll on moose, forcing them to near extinction, they are protected, but enforcement of the policy is weak and demand for traditional medicines derived from deer parts is high.^{[citation needed]} In 1978, the Regional Hunting Department transported 45 young moose to the center of Kamchatka. These moose were brought from Chukotka, home to the largest moose on the planet. Kamchatka now regularly is responsible for the largest trophy moose shot around the world each season. As it is a fertile environment for moose, with a milder climate, less snow, and an abundance of food, moose quickly bred and settled along the valley of the Kamchatka River and many surrounding regions. The population in the past 20 years has risen to over 2,900 animals.

The size of the moose varies. Following Bergmann's rule, population in the south (A. a. cameloides) usually grow smaller, while moose in the north and northeast (A. a. buturlini) can match the imposing sizes of the Alaskan moose (A. a. gigas) and are prized by trophy hunters.

New Zealand

In 1900, an attempt to introduce moose into the Hokitika area failed; then in 1910 ten moose (four bulls and six cows) were introduced into Fiordland. This area is considered a less than suitable habitat, and subsequent low numbers of sightings and kills have led to some presumption of this population's failure.^[142] The last proven sighting of a moose in New Zealand was in 1952.^[143] However, a moose antler was found in 1972, and DNA tests showed that hair collected in 2002 was from a moose. There has been extensive searching, and while automated cameras failed to capture photographs, evidence was seen of bedding spots, browsing, and antler marks.^[144]

Subspecies

History

European rock drawings and cave paintings reveal that moose have been hunted since the Stone Age. Excavations in Alby, Sweden, adjacent to the Stora Alvaret have yielded moose antlers in wooden hut remains from 6000 BCE, indicating some of the earliest moose hunting in northern Europe. In northern Scandinavia one can still find remains of trapping pits used for hunting moose. These pits, which can be up to 4 m × 7 m (13 ft × 23 ft) in area and 2 m (6 ft 7 in) deep, would have been camouflaged with branches and leaves^{[citation needed]}. They would have had steep sides lined with planks, making it impossible for the moose to escape once it fell in. The pits are normally found in large groups, crossing the moose's regular paths and stretching over several km. Remains of wooden fences designed to guide the animals toward the pits have been found in bogs and peat. In Norway, an early example of these trapping devices has been dated to around 3700 BC^{[citation needed]}. Trapping elk in pits is an extremely effective hunting method. As early as the 16th century the Norwegian government tried to restrict their use; nevertheless, the method was in use until the 19th century.

The earliest recorded description of the moose is in Julius Caesar's Commentarii de Bello Gallico, where it is described thus:

There are also [animals], which are called alces (moose). The shape of these, and the varied color of their skins, is much like roes, but in size they surpass them a little and are destitute of horns, and have legs without joints and ligatures; nor do they lie down for the purpose of rest, nor, if they have been thrown down by any accident, can they raise or lift themselves up. Trees serve as beds to them; they lean themselves against them, and thus reclining only slightly, they take their rest; when the huntsmen have discovered from the footsteps of these animals whither they are accustomed to betake themselves, they either undermine all the trees at the roots, or cut into them so far that the upper part of the trees may appear to be left standing. When they have leant upon them, according to their habit, they knock down by their weight the unsupported trees, and fall down themselves along with them.^[181]

In book 8, chapter 16 of Pliny the Elder's Natural History from 77 CE, the elk and an animal called achlis, which is presumably the same animal, are described thus:

... there is, also, the moose, which strongly resembles our steers, except that it is distinguished by the length of the ears and of the neck. There is also the achlis, which is produced in the land of Scandinavia; it has never been seen in this city, although we have had descriptions of it from many persons; it is not unlike the moose, but has no joints in the hind leg. Hence, it never lies down, but reclines against a tree while it sleeps; it can only be taken by previously cutting into the tree, and thus laying a trap for it, as otherwise, it would escape through its swiftness. Its upper lip is so extremely large, for which reason it is obliged to go backwards when grazing; otherwise, by moving onwards, the lip would get doubled up.^[182]

As food

Moose are hunted as a game species in many of the countries where they are found. Moose meat tastes, wrote Henry David Thoreau in The Maine Woods, "like tender beef, with perhaps more flavour; sometimes like veal". While the flesh has protein levels similar to those of other comparable red meats (e.g. beef, deer and wapiti), it has a low fat content, and the fat that is present consists of a higher proportion of polyunsaturated fats than saturated fats.^[183]

Dr. Valerius Geist, who emigrated to Canada from the Soviet Union, wrote in his 1999 book Moose: Behaviour, Ecology, Conservation:

In Sweden, no fall menu is without a mouthwatering moose dish. The Swedes fence their highways to reduce moose fatalities and design moose-proof cars. Sweden is less than half as large as the Canadian province of British Columbia, but the annual take of moose in Sweden—upward of 150,000—is twice that of the total moose harvest in North America.

Boosting moose populations in Alaska for hunting purposes is one of the reasons given for allowing aerial or airborne methods to remove wolves in designated areas, e.g., Craig Medred: "A kill of 124 wolves would thus translate to [the survival of] 1488 moose or 2976 caribou or some combination thereof".^[184] Some scientists believe that this artificial inflation of game populations is actually detrimental to both caribou and moose populations as well as the ecosystem as a whole. This is because studies have shown^{[citation needed]} that when these game populations are artificially boosted, it leads to both habitat destruction and a crash in these populations.^[185]

Vehicle collisions

The center of mass of a moose is above the hood of most passenger cars. In a collision, the impact crushes the front roof beams and individuals in the front seats.^[190] Collisions of this type are frequently fatal; seat belts and airbags offer little protection.^[191] In collisions with higher vehicles (such as trucks), most of the deformation is to the front of the vehicle and the passenger compartment is largely spared. Moose collisions have prompted the development of a vehicle test referred to as the "moose test" (Swedish: Älgtest, German: Elchtest). A Massachusetts study found that moose–vehicular collisions had a very high human fatality rate and that such collisions caused the death of 3% of the Massachusetts moose population annually.^[192]

Moose warning signs are used on roads in regions where there is a danger of collision with the animal. The triangular warning signs common in Sweden, Norway, and Finland have become coveted souvenirs among tourists traveling in these countries, causing road authorities so much expense that the moose signs have been replaced with imageless generic warning signs in some regions.^[193]

In Ontario, Canada, an estimated 265 moose die each year as a result of collision with trains (as of 2019^[update]). Moose–train collisions were more frequent in winters with above-average snowfall.^[194] In January 2008, the Norwegian newspaper Aftenposten estimated that some 13,000 moose had died in collisions with Norwegian trains since 2000. The state agency in charge of railroad infrastructure (Jernbaneverket) plans to spend 80 million Norwegian kroner to reduce collision rate in the future by fencing the railways, clearing vegetation from near the tracks, and providing alternative snow-free feeding places for the animals elsewhere.^[195]

In the Canadian province of New Brunswick, collisions between automobiles and moose are frequent enough that all new highways have fences to prevent moose from accessing the road, as has long been done in Finland, Norway, and Sweden. A demonstration project, Highway 7 between Fredericton and Saint John, which has one of the highest frequencies of moose collisions in the province, did not have these fences until 2008, although it was and continues to be extremely well signed.^[196][197] Newfoundland and Labrador recommended that motorists use caution between dusk and dawn because that is when moose are most active and most difficult to see, increasing the risk of collisions.^[198] Local moose sightings are often reported on radio stations so that motorists can take care while driving in particular areas. An electronic "moose detection system" was installed on two sections of the Trans-Canada Highway in Newfoundland in 2011, but the system proved unreliable and was removed in 2015.^[199] As of 2024^[update], the moose population in Newfoundland was increasing along with the number of road accidents.^[200]

In Sweden, a road will not be fenced unless it experiences at least one moose accident per km per year.^[201]

In eastern Germany, where the scarce population is slowly increasing, there were two road accidents involving moose since 2000.^[135]

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  Canadian road sign.
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  Finnish road sign.
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  Norwegian road sign.
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  Swedish road sign.
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  Warning sign in Alaska where trees and brush are trimmed along high moose crossing areas so that moose can be seen as they approach the road.
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  Moose (A. a. gigas) crossing a road in Alaska.

Domestication

Domestication of moose was investigated in the Soviet Union before World War II. Early experiments were inconclusive, but with the creation of a moose farm at Pechora-Ilych Nature Reserve in 1949, a small-scale moose domestication program was started, involving attempts at selective breeding of animals on the basis of their behavioural characteristics. Since 1963, the program has continued at Kostroma Moose Farm, which had a herd of 33 tame moose as of 2003. Although at this stage the farm is not expected to be a profit-making enterprise, it obtains some income from the sale of moose milk and from visiting tourist groups. Its main value, however, is seen in the opportunities it offers for the research in the physiology and behavior of the moose, as well as in the insights it provides into the general principles of animal domestication.

In Sweden, there was a debate in the late 18th century about the national value of using the moose as a domestic animal. Among other things, the moose was proposed to be used in postal distribution, and there was a suggestion to develop a moose-mounted cavalry. Such proposals remained unimplemented, mainly because the extensive hunting for moose that was deregulated in the 1790s nearly drove it to extinction. While there have been documented cases of individual moose being used for riding and/or pulling carts and sleds, Björklöf concludes no wide-scale usage has occurred outside fairy tales.^[202]

Heraldry

As one of the Canadian national symbols, the moose occurs on several Canadian coats of arms, including Newfoundland and Labrador,^[203] and Ontario.^[204] Moose is also a common coat of arms in Europe as well; for example, in Finland, it appears on the coats of arms of Hirvensalmi and Mäntsälä municipalities.^[205] The Seal of Michigan features a moose.^[206]