Themuskox (Ovibos moschatus)[a][b] is ahoofed mammal of the familyBovidae.[8] Native to theArctic, it is noted for its thick coat and for the strong odor emitted by males during theseasonal rut, from which its name derives. Thismusky odor has the effect of attracting females duringmating season. ItsInuktitut name "umingmak" translates to "the bearded one".[9]
ItsWoods Cree names "mâthi-môs" and "mâthi-mostos" translate to "ugly moose" and "ugly bison", respectively.[10] In historic times, muskoxen primarily lived inGreenland and theCanadian Arctic of theNorthwest Territories andNunavut.[11] They were formerly present in Eurasia, with their youngest natural records in the region dating to around 2,700 years ago,[12] with reintroduced populations in theAmerican state ofAlaska, theCanadian territory ofYukon, andSiberia, and an introduced population inNorway, part of which emigrated toSweden, where a small population now lives.
The muskox is in the subtribeOvibovina (or tribe Ovibovini) in the tribeCaprini (or subfamily Caprinae) of the subfamilyAntilopinae in the family Bovidae. It is therefore more closely related tosheep andgoats than tooxen; it is placed in its own genus,Ovibos (Latin: "sheep-ox"). It is one of the two largestextant members of the caprines, along with the similarly sizedTakinBudorcas.[13] While the takin and muskox were once considered possibly closely related, the takin lacks common ovibovine features, such as the muskox's specialized horn morphology, and genetic analysis shows that their lineages actually separated early in caprine evolution. Instead, the muskox's closest living relatives appear to be thegorals of the genusNaemorhedus, nowadays common in many countries of central and east Asia. The vague similarity between takin and muskox is therefore an example ofconvergent evolution.[14]
The modern muskox is the last member of a line of ovibovines that first evolved in temperate regions of Asia and adapted to a coldtundra environment late in its evolutionary history. Muskox ancestors with sheep-like high-positioned horns (horn cores being mostly over the plane of thefrontal bones, rather than below them as in modern muskoxen) first left the temperate forests for the developing grasslands ofCentral Asia during thePliocene, expanding intoSiberia and the rest of northernEurasia. Later migration waves of Asian ungulates that included high-horned muskoxen reachedEurope andNorth America during the first half of thePleistocene. The first well known muskox, the "shrub-ox"Euceratherium, crossed to North America over an early version of theBering Land Bridge two million years ago and prospered in theAmerican southwest andMexico.Euceratherium was larger yet more lightly built than modern muskoxen, resembling a giant sheep with massive horns, and preferred hilly grasslands.
A genus with intermediate horns,Soergelia, inhabited Eurasia in the early Pleistocene, fromSpain to Siberia, and crossed to North America during theIrvingtonian (1.8 million years to 240,000 years ago), soon afterEuceratherium. UnlikeEuceratherium, which survived in America until the Pleistocene-Holoceneextinction event,Soergelia was a lowland dweller which disappeared fairly early, displaced by more advanced ungulates, such as the "giant muskox"Praeovibos (literally "beforeOvibos"). The low-hornedPraeovibos was present in Europe and theMediterranean 1.5 million years ago, colonizedAlaska and theYukon one million years ago and disappeared half a million years ago.Praeovibos was a highly adaptable animal apparently associated with cold tundra (reindeer) and temperate woodland (red deer) faunas alike. During theMindel glaciation 500,000 years ago,Praeovibos was present in theKolyma river area in eastern Siberia in association with manyIce Agemegafauna that would later coexist withOvibos, in the Kolyma itself and elsewhere, includingwild horses, reindeer,woolly mammoth andstag-moose. It is debated, however, ifPraeovibos was directly ancestral toOvibos, or both genera descended from a common ancestor, since the two occurred together during the middle Pleistocene. Defenders of ancestry fromPraeovibos have proposed thatPraeovibos evolved intoOvibos in one region during a period of isolation and expanded later, replacing the remaining populations ofPraeovibos.[14]
Two morePraeovibos-like genera were named in America in the 19th century,Bootherium andSymbos, which are now identified as the male and female forms of a single,sexually dimorphic species, the "woodland muskox",Bootherium bombifrons.Bootherium inhabited open woodland areas of North America during the late Pleistocene, from Alaska toTexas and maybe even Mexico, but was most common in theSouthern United States, whileOvibos replaced it in the tundra-steppe to the north, immediately south of theLaurentian ice sheet.[14][15]
ModernOvibos appeared inGermany almost one million years ago and was common in the region through the Pleistocene. By the Mindel, muskoxen had also reached theBritish Isles. Both Germany and Britain were just south of theScandinavian ice sheet and covered in tundra during cold periods, but Pleistocene muskoxen are also rarely recorded in more benign and wooded areas to the south likeFrance andGreen Spain, where they coexisted with temperate ungulates likered deer andaurochs. Likewise, the muskox is known to have survived in Britain during warminterglacial periods.[14]
Today's muskoxen are descended from others believed to have migrated fromSiberia toNorth America between 200,000[16] and 90,000 years ago,[17] having previously occupied Alaska (at the time united to Siberia and isolated periodically from the rest of North America by the union of the Laurentide andCordilleran Ice Sheets during colder periods) between 250,000 and 150,000 years ago. After migrating south during one of the warmer periods of theIllinoian glaciation, non-Alaskan American muskoxen would be isolated from the rest in the colder periods. The muskox was already present in its current stronghold ofBanks Island 34,000 years ago, but the existence of other ice-free areas in theCanadian Arctic Archipelago at the time is disputed.[14]
Fossil DNA evidence suggests that muskoxen were not only more geographically widespread during the Pleistocene, but also moregenetically diverse.[20] During that time, other populations of muskoxen lived across the Arctic, from theUral Mountains to Greenland. By contrast, the current genetic makeup of the species is more homogenous. Climate fluctuation may have affected this shift in genetic diversity: research indicates colder periods in Earth's history are correlated with more diversity, and warmer periods with more homogeneity.[19] Muskox populations survived into the Holocene in Siberia, with their youngest records in the region being from theTaymyr Peninsula, dating to around 2,700 years ago (~700 BC).[12]
Both male and female muskoxen have long, curvedhorns. Muskoxen stand 1.1 to 1.5 m (3 ft 7 in to 4 ft 11 in) high atwithers, with females measuring 135 to 200 cm (4 ft 5 in to 6 ft 7 in) in length, and the larger males 200 to 250 cm (6 ft 7 in to 8 ft 2 in). The small tail, often concealed under a layer of fur, measures only 10 cm (3.9 in) long. Adults, on average, weigh 285 kg (628 lb), but can range from 180 to 410 kg (400 to 900 lb).[13][21] The thick coat and large head suggest a larger animal than the muskox truly is; the bison, to which the muskox is often compared, can weigh up to twice as much.[22] However, heavy zoo-kept specimens have weighed up to 650 kg (1,430 lb).[7] Their coat, a mix of black, gray and brown, includes longguard hairs that almost reach the ground. Rare "white muskoxen" have been spotted in theQueen Maud Gulf Bird Sanctuary.[23] Muskoxen are occasionally semi-domesticated for wool, and rarely for meat and milk. The U.S. state ofAlaska has several muskoxen farms specifically aimed at wool harvesting.[24][25] Thewool, calledqiviut, is highly prized for its softness, length, and insulation value. Prices for yarn range between $1.5 and $3/g ($40 and $80/oz).[26][27][28]
A muskox can reach speeds of up to 60 km/h (37 mph).[29] Their life expectancy is between 12 and 20 years.
During thePleistocene period, muskoxen were much more widespread. Fossil evidence shows that they lived across the Siberian and North American Arctic, from theUrals toGreenland.[19] The ancestors of today's muskoxen came across theBering Land Bridge to North America between 200,000[16] and 90,000 years ago.[17] During theWisconsinan, modern muskox thrived in thetundra south of theLaurentide Ice Sheet, in what is now theMidwest, theAppalachians andVirginia, while distant relativesBootherium andEuceratherium lived in the forests of theSouthern United States and the western shrubland, respectively.[15] Though they were always less common than other Ice Age megafauna, muskox abundance peaked during theWürm II glaciation 20,000 years ago and declined afterwards, especially during thePleistocene/Holoceneextinction event, where its range was greatly reduced and only the populations in North America survived. The last known muskox population in Europe died out inSweden 9,000 years ago.[14] In Asia, muskox persisted until just 615-555 BCE inTumat,Sakha Republic.[30][failed verification]
Following the disappearance of theLaurentide Ice Sheet, the muskox gradually moved north across theCanadian Arctic Archipelago, arriving inGreenland fromEllesmere Island at about 350 AD, during the lateHolocene. Their arrival in northwestern Greenland probably occurred within a few hundred years of the arrival of theDorset andThule cultures in the present-dayQaanaaq area. Human predation around Qaanaaq may have restricted muskoxen from moving down the west coast, and instead kept them confined to the northeastern fringes of the island.[31]
There have been at least two domestication endeavours. In the 1950s, an American researcher and adventurer was able to capture muskox calves in Northern Canada for relocation to a property he prepared in Vermont.[38][39][40] One condition imposed by the Canadian government was that he was not allowed to kill adults defending their young. When nets and ropes proved useless, he and his crew herded family groups into open water, where calves were successfully separated from the adults. Once airfreighted to Montreal and trucked to Vermont, the young animals habituated to the temperate conditions. Although the calves thrived and grew to adulthood, parasite and disease resistance problems impaired the overall success of the effort. The surviving herd was eventually moved to a farm inPalmer, Alaska, where it has been successful since the mid-1950s.[41]
In 1913, workers building a railway overDovrefjell found two fossil muskox vertebrae. This led to the idea of introducing muskoxen to Norway from Greenland. The first release in the world was made onGurskøy outsideÅlesund in 1925–26. They were muskoxen caught by Norwegian seal-hunting boats in Greenland. The animals colonized the island, but eventually died out there. An attempt to introduce the muskox toSvalbard also failed. Seventeen animals were released in 1929 byAdventfjorden onWest Spitsbergen. In 1940, the herd numbered 50, but in the 1970s, the whole herd disappeared. In September 1932, polar researcherAdolf Hoel conducted another experiment, importing 10 muskoxen to Dovrefjell. This herd survived untilWorld War II, when they were hunted and exterminated. In 1947 and later, new animals were released. A small group of muskoxen from Dovrefjell migrated across the national border to Sweden in 1971 and established themselves inHärjedalen, whereby a Swedish herd was established.[citation needed]
The Norwegian population on Dovrefjell is managed over an area of 340 km2 (130 sq mi) and in the summer of 2012 consisted of approximately 300 animals. Since 1999, the population has mostly been increasing, but it suffered a measles outbreak in the summer of 2004 that killed 29. Some animals are also occasionally killed as a result of train collisions on theDovre Railway. The population is divided into flocks in theNystuguhø [no] area,Kolla [no] area andHjerkinn. In the summer they move down towardsDriva, where there are lush grass pastures.
Although the muskox belongs to the dry Arctic grassland, it seems to do well on Dovrefjell. However, the pastures are marginal, with little grass available in winter (the muskox eats only plants, not lichen as reindeer do), and over time,inbreeding depression is expected in such a small population which originated from only a few introduced animals.
Muskoxen were introduced toSvalbard in 1925–26 and 1929, but this population died out in the 1970s.[42] They were also introduced inIceland around 1930 but did not survive.[43]
InRussia, animals imported from Banks and Nunivak were released in theTaymyr Peninsula in 1974 and 1975, and some from Nunivak were released inWrangel Island in 1975. Both locations are north of theArctic Circle. By 2019 the population on Wrangel Island was about 1100,[44] and the Taymyr Peninsula, about 11,000–14,000.[45] A few muskoxen herds migrated from the Taymyr Peninsula far to the south to thePutorana Plateau.[44] Once established, these populations have been, in turn, used as sources for further reintroductions in Siberia between 1996 and 2010.[46] One of the last of these actions was the release of six animals within thePleistocene Park project area in theKolyma River in 2010, where a team of Russian scientists led bySergey Zimov aims to prove that muskoxen, along with otherPleistocene megafauna that survived into the earlyHolocene in northern Siberia,[47] did not disappear from the region due to climate change, but because of human hunting.[48]
Ancient muskox remains have never been found ineastern Canada, although the ecological conditions in the northernLabrador Peninsula are suitable for them. In 1967, 14 animals were captured nearEureka onEllesmere Island by the Institute for Northern Agricultural Research (INAR) and brought to a farm in Old Fort ChimoKuujjuaq, northern Quebec, for domestication to provide a local cottage industry based onqiviut, a fine natural fiber. The animals thrived and theqiviut industry showed early success with the training of Inuit knitters and marketing, but it soon became clear that the Quebec government had never intended that the muskoxen be domestic, but had used INAR to capture muskoxen to provide a wild population for hunting[citation needed]. Government officials demanded that INAR leave Quebec and the farm be closed. Subsequently, 54 animals from the farm were released in three places in northern Quebec between 1973 and 1983, and the remaining were ceded to localzoos. Between 1983 and 1986, the released animals increased from 148 to 290, at a rate of 25% per year, and by 2003, an estimated 1,400 muskoxen were in Quebec. Additionally, 112 adults and 25 calves were counted in the nearbyDiana Island in 2005, having arrived there by their own means from the mainland.Vagrant adults are sometimes spotted inLabrador, though no herds have been observed in the region.[49]
During the summer, muskoxen live in wet areas, such as river valleys, moving to higher elevations in the winter to avoid deep snow. Muskoxen will eat grasses,arctic willows, woody plants, lichens(above lichens are excluded from the menu), and mosses. When food is abundant, they prefer succulent and nutritious grasses in an area. Willows are the most commonly eaten plants in the winter. Muskoxen require a high threshold of fat reserves in order to conceive, which reflects their conservative breeding strategy. Winter ranges typically have shallow snow to reduce the energy costs of digging through snow to reach forage.[1] The primary predators of muskoxen arearctic wolves, which may account for up to half of all mortality for the species. Other occasional predators, likely mainly predators of calves or infirm adults, can includegrizzly bears andpolar bears[7] andwolverines.[dubious –discuss][citation needed]
Muskox areheterothermic mammals, meaning they have the ability to shut off thermal regulation in some parts of their body, like their lower limbs.[50] Maintaining the lower limbs at a cooler temperature than the rest of their body helps reduce the loss of body heat from their extremities. Muskox display the unique characteristic of havinghemoglobin that is three times less temperature sensitive than human hemoglobin.[50] This temperature insensitivity allows the muskox's hemoglobin to have a heightened oxygen affinity in an extremely cold environment and continue to diffuse high amounts of oxygen into its cold tissues.[51]
Nunivak Island, Alaskan muskoxen in the 1930s, shown here in defensive formation
Muskoxen live in herds of 12–24 in the winter and 8–20 in the summer when dominant bulls expel other males from the herd.[52] They do not hold territories, but they do mark their trails withpreorbital glands.[53] Male and female muskoxen have separate age-based hierarchies, with mature oxen being dominant over juveniles.[52] Dominant oxen tend to get access to the best resources[7] and will displace subordinates from patches of grass during the winter.[52] Muskox bulls assert their dominance in many different ways. One is a "rush and butt", in which a dominant bull rushes a subordinate from the side with its horns, and will warn the subordinate so it can have a chance to get away.[54] Bulls will also roar, swing their heads, and paw the ground.[7] Dominant bulls sometimes treat subordinate bulls like cows. A dominant bull will tap a subordinate with its foreleg, something they do to cows during mating.[55] Dominant bulls will also mock copulate subordinates and sniff their genitals.[55] A subordinate bull can challenge his status by charging a dominant bull.[56]
The mating (or "rutting") season of the muskoxen begins in late June or early July. During this time, dominant bulls will fight others out of the herds and establish harems of usually six or seven cows and their offspring. Fighting bulls will first rub their preorbital glands against their legs while bellowing loudly, and then display their horns.[56] The bulls then back up about 20 m (66 ft), lower their heads, and charge into each other, and will keep doing so until one bull gives up.[54] Subordinate and elderly bulls will leave the herds to form bachelor groups or become solitary.[7] However, when danger is present, the outside bulls can return to the herd for protection.[57] Dominant bulls will prevent cows from leaving their harems.[7] During mating, a bull will tap an estrous cow with his foreleg to calm her down and make her more receptive to his advances.[55] The herds reassemble when summer ends.[57]
While the bulls are more aggressive during the rutting season and lead their groups, the females take charge during gestation.[7] Pregnant females are aggressive and decide what distance the herd travels in a day and where they will bed for the night.[58] The herds move more often when cows are lactating, to let them get enough food to nurse their offspring.[58] Cows have an eight- to nine-month gestation period, with calving occurring from April to June. Cows do not calve every year. When winters are severe, cows will not go into estrus and thus not calve the next year. When calving, cows stay in the herd for protection. Muskox areprecocial, and calves can keep up with the herd within just a few hours after birth. The calves are welcomed into the herd and nursed for the first two months.[7] After that, a calf then begins eating vegetation and nurses only occasionally. Cows communicate with their calves through braying. The calf's bond with its mother weakens after two years.
Muskoxen have a distinctive defensive behavior: when the herd is threatened, the adults will face outward to form a stationary ring or semicircle around the calves.[59] The bulls are usually the front line for defense against predators, with the cows and juveniles gathering close to them.[7] Bulls determine the defensive formation during rutting, while the cows decide the rest of the year.[57]
The preorbital gland secretion of muskoxen has a "light, sweetish, ethereal" odor.[9] Analysis of preorbital gland secretion extract showed the presence ofcholesterol (which is nonvolatile),benzaldehyde, a series of straight-chain saturatedγ-lactones ranging from C8H14O2 to C12H22O2 (with C10H18O2 being most abundant), and probably the monounsaturated γ-lactone C12H20O2.[9] The saturated γ-lactone series has an odor similar to that of the secretion.[9]
Muskoxen are not known to be aggressive. Fatal attacks are extremely rare, but humans who have come close and behaved aggressively have occasionally been attacked.[60]
On 22 July 1964, a 73-year-old man was killed in a muskox attack in Norway. The animal was later killed by local authorities.[61]
On 13 December 2022, a court services officer with theAlaska State Troopers was killed by a muskox nearNome, Alaska. The officer was trying to scare away a group of muskox near a dog kennel at his home when one of the animals attacked him.[62]
Historically, this species declined because of overhunting, but populations have recovered following enforcement of hunting regulations.[1] Management in the late 1900s was mostly conservative hunting quotas to foster recovery and recolonization from the historic declines.[1] The current world population of muskoxen is estimated at between 80,000[63] and 125,000,[34] with an estimated 47,000 living onBanks Island.[64]
In Greenland, there are no major threats. However, populations are often small in size and scattered; this makes them vulnerable to local fluctuations in climate. Most populations are within national parks, where they are protected from hunting.[1] Muskoxen occur in four of Greenland's protected areas, with indigenous populations inNortheast Greenland National Park and introduced populations inArnangarnup Qoorua Nature Reserve [d] andKangerlussuaq andManiitsoq Caribou Reserves [d]. In these areas, muskoxen receive full protection.[1]
Muskoxen are being domesticated for the production ofqiviut.
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