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The reindeer or caribou is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, boreal, and mountainous regions of Northern Europe, Siberia, and North America. It is the only representative of the genus Rangifer. More recent studies suggest the splitting of reindeer and caribou. "All caribou and reindeer throughout the world are considered to be the same species, but there are 7 subspecies."

Description
Names follow international convention before the recent revision (see below). Reindeer / caribou (Rangifer) vary in size from the smallest, the Svalbard reindeer (R. (t.) platyrhynchus), to the largest, Osborn's caribou (R. t. osborni). They also vary in coat color and antler architecture. The North American range of caribou extends from Alaska through the Yukon, the Northwest Territories and Nunavut throughout the tundra, taiga (boreal forest) and south through the Canadian Rocky Mountains. Of the eight subspecies classified by Harding (2022) into the Arctic caribou (R. arcticus), the migratory mainland barren-ground caribou of Arctic Alaska and Northern Canada (R. t. arcticus), summer in tundra and winter in taiga, a transitional forest zone between boreal forest and tundra; the nomadic Peary caribou (R. t. pearyi) lives in the polar desert of the high Arctic Archipelago and Grant's caribou (R. t. granti also called the Porcupine caribou) lives in the western end of the Alaska Peninsula and the adjacent islands; the other four subspecies, Osborn's caribou (R. t. osborni), Stone's caribou (R. t. stonei), the Rocky Mountain caribou (R. t. fortidens) and the Selkirk Mountains caribou (R. t. montanus) are all montane. The extinct insular Queen Charlotte Islands caribou (R. t. dawsoni), lived on Graham Island in Haida Gwaii (formerly known as the Queen Charlotte Islands). The boreal woodland caribou (R. t. caribou), lives in the boreal forest of northeastern Canada: the Labrador or Ungava caribou of northern Quebec and northern Labrador (R. t. caboti), and the Newfoundland caribou of Newfoundland (R. t. terranovae) have been found to be genetically in the woodland caribou lineage. and Geist, 1998). == Status ==
Status
About 25,000 mountain reindeer (R. t. tarandus) still live in the mountains of Norway, notably in Hardangervidda. In Sweden there are approximately 250,000 reindeer in herds managed by Sámi villages. Russia manages 19 herds of Siberian tundra reindeer (R. t. sibiricus) that total about 940,000. The Taimyr herd of Siberian tundra reindeer is the largest wild reindeer herd in the world, varying between 400,000 and 1,000,000; it is a metapopulation consisting of several subpopulations — some of which are phenotypically different — with different migration routes and calving areas. The Kamchatkan reindeer (R. t. phylarchus), a forest subspecies, formerly included reindeer west of the Sea of Okhotsk which, however, are indistinguishable genetically from the Jano-Indigirka, East Siberian taiga and Chukotka populations of R. t. sibiricus. Siberian tundra reindeer herds have been in decline but are stable or increasing since 2000. The New York Times reported in April 2018 of the disappearance of the only herd of southern mountain woodland caribou in the contiguous United States, with an expert calling it "functionally extinct" after the herd's size dwindled to a mere three animals. After the last individual, a female, was translocated to a wildlife rehabilitation center in Canada, caribou were considered extirpated from the contiguous United States. The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) classified both the Southern Mountain population DU9 (R. t. montanus) and the Central Mountain population DU8 (R. t. fortidens) as Endangered and the Northern Mountain population DU7 (R. t. osborni) as Threatened. Some species and subspecies are rare and three subspecies have already become extinct: the Queen Charlotte Islands caribou (R. t. dawsoni) from western Canada, the Sakhalin reindeer (R. t. setoni) from Sakhalin and the East Greenland caribou from eastern Greenland, although some authorities believe that the latter, R. t. eogroenlandicus Degerbøl, 1957, is a junior synonym of the Peary caribou. Historically, the range of the sedentary boreal woodland caribou covered more than half of Canada and into the northern states of the contiguous United States from Maine to Washington. Boreal woodland caribou have disappeared from most of their original southern range and were designated as Threatened in 2002 by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC). Environment and Climate Change Canada reported in 2011 that there were approximately 34,000 boreal woodland caribou in 51 ranges remaining in Canada (Environment Canada, 2011b), although those numbers included montane populations classified by Harding (2022) into subspecies of the Arctic caribou. Siberian tundra reindeer herds are also in decline, and Rangifer as a whole is considered to be Vulnerable by the International Union for Conservation of Nature (IUCN). == Naming ==
Naming
Charles Hamilton Smith is credited with the name Rangifer for the reindeer genus, which Albertus Magnus used in his , fol. Liber 22, Cap. 268: "Dicitur Rangyfer quasi ramifer". This word may go back to the Sámi word . thought that rangifer and tarandus were two separate animals. In any case, the tarandos name goes back to Aristotle and Theophrastus. The use of the terms reindeer and caribou for essentially the same animal can cause confusion, but the International Union for Conservation of Nature clearly delineates the issue: "Reindeer is the European name for the species of Rangifer, while in North America, Rangifer species are known as Caribou." According to the US Food and Drug Administration, in North American English the animal is known as caribou if wild and reindeer if domesticated. The word reindeer is an anglicized version of the Old Norse words ("reindeer") and ("animal") and has nothing to do with reins. The word caribou comes through French, from the Mi'kmaq , meaning "snow shoveler", and refers to its habit of pawing through the snow for food. Because of its importance to many cultures, Rangifer and some of its species and subspecies have names in many languages. Inuvialuit of the western Canadian Arctic and Inuit of the eastern Canadian Arctic, who speak different dialects of the Inuit languages, both call the barren-ground caribou . The Wekʼèezhìi (Tłı̨chǫ) people, a Dene (Athapascan) group, call the Arctic caribou and the boreal woodland caribou . The Gwichʼin (also a Dene group) have over 24 distinct caribou-related words. Reindeer are also called by the Greenlandic Inuit and , sometimes , by the Icelanders. == Evolution ==
Evolution
The "glacial-interglacial cycles of the upper Pleistocene had a major influence on the evolution" of Rangifer species and other Arctic and sub-Arctic species. Isolation of tundra-adapted species Rangifer in Last Glacial Maximum refugia during the last glacial – the Wisconsin glaciation in North America and the Weichselian glaciation in Eurasia – shaped "intraspecific genetic variability" particularly between the North American and Eurasian parts of the Arctic. The Eurasian clade of Odocoileinae (Capreolini, Hydropotini and Alcini) split from the New World tribes of Capreolinae (Odocoileini and Rangiferini) in the Late Miocene, 8.7–9.6 million years ago. Rangifer "evolved as a mountain deer, ...exploiting the subalpine and alpine meadows...". As well, many genes, including those for vitamin D metabolism, fat metabolism, retinal development, circadian rhythm, and tolerance to cold temperatures, are found in tundra caribou that are lacking or rudimentary in forest types. For this reason, forest-adapted reindeer and caribou could not survive in tundra or polar deserts. The oldest undoubted Rangifer fossil is from Omsk, Russia, dated to 2.1-1.8 million years before present (BP). The oldest North American Rangifer fossil is from the Yukon, 1.6 million years BP. A fossil skull fragment from Süßenborn, Germany, R. arcticus stadelmanni, (which is probably misnamed) with "rather thin and cylinder-shaped" antlers, dates to the Middle Pleistocene (Günz) Period, 680,000-620,000 BP. Rangifer fossils become increasingly frequent in circumpolar deposits beginning with the Riss glaciations, the second youngest of the Pleistocene Epoch, roughly 300,000–130,000 BP. By the 4-Würm period (110,000–70,000 to 12,000–10,000 BP), its European range was extensive, supplying a major food source for prehistoric Europeans. North American fossils outside of Beringia that predate the Last Glacial Maximum (LGM) are of Rancholabrean age (240,000–11,000 years BP) and occur along the fringes of the Rocky Mountain and Laurentide ice sheets as far south as northern Alabama, USA; and in Sangamonian deposits (~100,000 years BP) from western Canada. A R. t. pearyi-sized caribou occupied Greenland before and after the LGM and persisted in a relict enclave in northeastern Greenland until it went extinct about 1900 (see discussion of R. t. eogroenlandicus below). Archaeological excavations showed that larger barren-ground-sized caribou appeared in western Greenland about 4,000 years ago. Valerius Geist (1998) Finnish forest reindeer (R. t. fennicus) likely evolved from Cervus [Rangifer] guettardi Desmarest, 1822, a reindeer that adapted to forest habitats in Eastern Europe as forests expanded during an interglacial period before the LGM (the Würmian or Weichselian glaciation);. in a second immigration 19,000–20,000 years ago when the LGM turned its forest habitats into tundra, while fennicus survived in isolation in southwestern Europe. At that time, modern tundra caribou had not even evolved. Woodland caribou are likely more related to extinct North American forest caribou than to barren-ground caribou. For example, the extinct caribou Torontoceros [Rangifer] hypogaeus, had features (robust and short pedicles, smooth antler surface, and high position of second tine) that relate it to forest caribou. Humans started hunting reindeer in both the Mesolithic and Neolithic Periods, and humans are today the main predator in many areas. Norway and Greenland have unbroken traditions of hunting wild reindeer from the Last Glacial Period until the present day. In the non-forested mountains of central Norway, such as Jotunheimen, it is still possible to find remains of stone-built trapping pits, guiding fences and bow rests, built especially for hunting reindeer. These can, with some certainty, be dated to the Migration Period, although it is not unlikely that they have been in use since the Stone Age. Cave paintings by ancient Europeans include both tundra and forest types of reindeer. == Taxonomy ==
Taxonomy
Naming and research on museum collections Carl Linnaeus in 1758 named the Eurasian tundra species Cervus tarandus, the genus Rangifer being credited to Smith, 1827. The low bez tines help the wide flat brow tines dig craters in the hard-packed tundra snow for forage, for which reason brow tines are often called "shovels" in North America and "ice tines" in Europe. The differences in antler architecture reflect fundamental differences in ecology and behavior, and in turn deep divisions in ancestry that were not apparent to the early taxonomists. Similarly, working on museum collections where skins were often faded and in poor states of preservation, early taxonomists could not readily perceive differences in coat patterns that are consistent within a subspecies, but variable among them. Geist calls these "nuptial" characteristics: sexually selected characters that are highly conserved and diagnostic among subspecies. Reclassification In the mid-20th century, as definitions of "species" evolved, mammalogists in Europe and North America made all Rangifer species conspecific with R. tarandus, and synonymized most of the subspecies. Alexander William Francis Banfield's often-cited A Revision of the Reindeer and Caribou, Genus Rangifer (1961), eliminated R. t. caboti (the Labrador caribou), R. t. osborni (Osborn's caribou — from British Columbia) and R. t. terranovae (the Newfoundland caribou) as invalid and included only barren-ground caribou, renamed as R. t. groenlandicus (formerly R. arcticus) and woodland caribou as R. t. caribou. However, Banfield made multiple errors, eliciting a scathing review by Ian McTaggart-Cowan in 1962. Most authorities continued to consider all or most subspecies valid; some were quite distinct. In his chapter in the authoritative 2005 reference work Mammal Species of the World, However, since 1991, many genetic studies have revealed deep divergence between modern tundra reindeer and woodland caribou. Geist (2007) and others continued arguing that the woodland caribou was incorrectly classified, noting that "true woodland caribou, the uniformly dark, small-maned type with the frontally emphasized, flat-beamed antlers", is "scattered thinly along the southern rim of North American caribou distribution". He affirms that the "true woodland caribou is very rare, in very great difficulties and requires the most urgent of attention." Ecotypes In 2011, noting that the former classifications of Rangifer tarandus, either with prevailing taxonomy on subspecies, designations based on ecotypes, or natural population groupings, failed to capture "the variability of caribou across their range in Canada" needed for effective subspecies conservation and management, COSEWIC developed Designatable Unit (DU) attribution, The 12 designatable units for caribou in Canada (that is, excluding Alaska and Greenland) based on ecology, behavior and, importantly, genetics (but excluding morphology and archaeology) essentially followed the previously named subspecies distributions, without naming them as such, plus some ecotypes. Ecotypes are not phylogenetically based and cannot substitute for taxonomy. Genetic, molecular, and archaeological evidence Meanwhile, genetic data continued to accumulate, revealing sufficiently deep divisions to easily separate Rangifer back into six previously named species and to resurrect several previously named subspecies. Molecular data showed that the Greenland caribou (R. t. groenlandicus) and the Svalbard reindeer (R. t. platyrhynchus), although not closely related to each other, were the most genetically divergent among Rangifer clades; Meanwhile, archaeological evidence was accumulating that Eurasian forest reindeer descended from an extinct forest-adapted reindeer and not from tundra reindeer (see Evolution above); since they do not share a direct common ancestor, they cannot be conspecific. Similarly, woodland caribou diverged from the ancestors of Arctic caribou before modern barren-ground caribou had evolved, and were more likely related to extinct North American forest reindeer (see Evolution above). Lacking a direct shared ancestor, barren-ground and woodland caribou cannot be conspecific. Molecular data also revealed that the four western Canadian montane ecotypes are not woodland caribou: they share a common ancestor with modern barren-ground caribou (tundra reindeer), but distantly, having diverged more than 60,000 years ago — before the modern ecotypes had evolved their cold- and darkness-adapted physiologies and mass-migration and aggregation behaviors (see Evolution above). Before Banfield (1961), taxonomists using cranial, dental and skeletal measurements had unequivocally allied these western montane ecotypes with barren-ground caribou, naming them (as in Osgood 1909 Murie, 1935 and Anderson 1946, among others) R. t. stonei, R. t. montanus, R. t. fortidens and R. t. osborni, respectively, Neither one of these clades has yet been formally described or named. Jenkins et al. (2012) said that "[Baffin Island] caribou are unique compared to other Barrenground herds, as they do not overwinter in forested habitat, nor do all caribou undertake long seasonal migrations to calving areas." It also shares a mtDNA haplotype with Labrador caribou, in the North American lineage (i.e., woodland caribou). Røed et al. (1991) had noted: Among Baffin Island caribou the TFL2 allele was the most common allele (p=0.521), while this allele was absent, or present in very low frequencies, in other caribou populations (Table 1), including the Canadian barren-ground caribou from the Beverly herd. A large genetic difference between Baffin Island caribou and the Beverly herd was also indicated by eight alleles found in the Beverly herd which were absent from the Baffin Island samples. Jenkins et al. (2018) also reported genetic distinctiveness of Baffin Island caribou from all other barren-ground caribou; its genetic signature was not found on the mainland or on other islands; nor were Beverly herd (the nearest mainly barren-ground caribou) alleles present in Baffin Island caribou, evidence of reproductive isolation. These advances in Rangifer genetics were brought together with previous morphological-based descriptions, ecology, behavior and archaeology to propose a new revision of the genus. == Species and subspecies ==
Species and subspecies
Abbreviations: • AMNH: American Museum of Natural HistoryBCPM: British Columbia Provincial Museum (= RBCM the Royal British Columbia Museum) • NHMUK: British Museum (Natural History) (originally the BMNH) • DMNH: Denver Museum of Natural HistoryMCZ: Museum of Comparative ZoologyMSI: Museum of the Smithsonian InstitutionNMC: National Museum of Canada (originally the CGS Canadian Geological Survey Museum, now the CMN Canadian Museum of Nature) • NR: Naturhistoriska RiksmuseetRSMNH: Royal Swedish Museum of Natural HistoryUSNM,: United States National MuseumZMASL: Zoological Museum of the Zoological Institute of the Russian Academy of Sciences (formerly the Zoological Museum of the Academy of Sciences), Leningrad The table above includes, as per the recent revision, R. t. caboti (the Labrador caribou (the Eastern Migratory population DU4)), and R. t. terranovae (the Newfoundland caribou (the Newfoundland population DU5)), which molecular analyses have shown to be of North American (i.e., woodland caribou) lineage; seem short of a taxonomic authority. illustrated and claimed to have seen a male specimen ("head of perfect horns...") from Greenland and said that a Captain Craycott had brought a live pair from Greenland to England in 1738. He named it Capra groenlandicus, Greenland reindeer. Linnaeus, in the 12th edition of Systema naturae, gave grœnlandicus as a synonym for Cervus tarandus. Borowski disagreed (and again changed the spelling), saying Cervus grönlandicus was morphologically distinct from Eurasian tundra reindeer. Baird placed it under the genus Rangifer as R. grœnlandicus. It went back and forth as a full species or subspecies of the barren-ground caribou (R. arcticus) or a subspecies of the tundra reindeer (R. tarandus), but always as the Greenland reindeer / caribou. Taxonomists consistently documented morphological differences between Greenland and other caribou / reindeer in cranial measurements, dentition, antler architecture, etc. Then Banfield (1961) R. arctus. However, because genetic data shows the Greenland caribou to be the most distantly related of any caribou to all the others (genetic distance, FST = 44%, by extending to them the name "granti". Valerius Geist (1998), in the only error in his whole illustrious career, re-analyzed Banfield's data with additional specimens found in an unpublished report he cites as "Skal, 1982", but was "not able to find diagnostic features that could segregate this form from the western barren ground type." But Skal 1982 had included specimens from the eastern end of the Alaska Peninsula and the Kenai Peninsula, the range of the larger Stone's caribou. Later, geneticists comparing barren-ground caribou of Alaska with those of mainland Canada found little difference and they all became the former R. t. groenlandicus (now R. t. arcticus). R. t. granti was lost in the oblivion of invalid taxonomy until Alaskan researchers sampled some small, pale caribou from the western end of the Alaska Peninsula, their range enclosing the type locality designated by Allen (1902) and found them to be genetically distinct from all other caribou in Alaska. Thus, granti was rediscovered, its range restricted to that originally described. Stone's caribou (R. t. stonei), It was placed under R. arcticus as a subspecies, North American examples of this are the Torngat Mountain population DU10, an ecotype of R. t. caboti; a recently discovered and unnamed clade between the Mackenzie River and Great Bear Lake of Beringian-Eurasian lineage, an ecotype of R. t. osborni; the Atlantic-Gaspésie population DU11, an eastern montane ecotype of the boreal woodland caribou (R. t. caribou); the Baffin Island caribou, an ecotype of the barren-ground caribou (R. t. arcticus); The last three of these likely qualify as subspecies, but they have not yet been formally described or named. == Physical characteristics ==
Physical characteristics
Naming in this and following sections follows the taxonomy in the authoritative 2011 reference work Handbook of the Mammals of the World Vol. 2: Hoofed Mammals. Androgens play an essential role in the antler formation of cervids. The antlerogenic genes in reindeer have more sensitivity to androgens in comparison with other cervids. There is considerable variation among species and subspecies in the size of the antlers (e.g., they are rather small and spindly in the northernmost species and subspecies), The number of points on male reindeer increases from birth to 5 years of age and remains relatively constant from then on. While antlers of male woodland caribou are typically smaller than those of male barren-ground caribou, they can be over across. They are flattened in cross-section, compact and relatively dense. The antlers' main beams begin at the brow "extending posterior over the shoulders and bowing so that the tips point forward. The prominent, palmate brow tines extend forward, over the face." The antlers typically have two separate groups of points, lower and upper. Antlers begin to grow on male reindeer in March or April and on female reindeer in May or June. This process is called antlerogenesis. Antlers grow very quickly every year on the bulls. As the antlers grow, they are covered in thick velvet, filled with blood vessels and spongy in texture. The antler velvet of the barren-ground caribou and the boreal woodland caribou is dark chocolate brown. The velvet that covers growing antlers is a highly vascularised skin. This velvet is dark brown on woodland or barren-ground caribou and slate-grey on Peary caribou and the Dolphin-Union caribou herd. Velvet lumps in March can develop into a rack measuring more than a in length by August. When the antler growth is fully grown and hardened, the velvet is shed or rubbed off. To Inuit, for whom the caribou is a "culturally important keystone species", the months are named after landmarks in the caribou life cycle. For example, amiraijaut in the Igloolik region is "when velvet falls off caribou antlers." In describing woodland caribou, which have a harem-defense mating system, SARA wrote, "During the rut, males engage in frequent and furious sparring battles with their antlers. Large males with large antlers do most of the mating." Reindeer continue to migrate until the bulls have spent their back fat. By contrast, barren-ground caribou males tend individual females and their fights are brief and much less intense; consequently, their antlers are long, and thin, round in cross-section and less branched and are designed more for show (or sexual attraction) than fighting. In late autumn or early winter after the rut, male reindeer lose their antlers, growing a new pair the next summer with a larger rack than the previous year. Female reindeer keep their antlers until they calve. In the Scandinavian and Arctic Circle populations, old bulls' antlers fall off in late December, young bulls' antlers fall off in the early spring, and cows' antlers fall off in the summer. When male reindeer shed their antlers in early to mid-winter, the antlered cows acquire the highest ranks in the feeding hierarchy, gaining access to the best forage areas. These cows are healthier than those without antlers. Calves whose mothers do not have antlers are more prone to disease and have a significantly higher mortality. caribou (tuktu) antlers According to the Igloolik Oral History Project (IOHP), "Caribou antlers provided the Inuit with a myriad of implements, from snow knives and shovels to drying racks and seal-hunting tools. A complex set of terms describes each part of the antler and relates it to its various uses". Pelt The color of the fur varies considerably, both between individuals and depending on season and species. Northern populations, which usually are relatively small, are whiter, while southern populations, which typically are relatively large, are darker. This can be seen well in North America, where the northernmost subspecies, the Peary caribou, is the whitest and smallest subspecies of the continent, while the Selkirk Mountains caribou (Southern Mountain population DU9) only exceeded in size by Osborn's caribou (Northern Mountain population DU7). Fur is the primary insulation factor that allows reindeer to regulate their core body temperature in relation to their environment, the thermogradient, even if the temperature rises to . In 1913, Dugmore noted how the woodland caribou swim so high out of the water, unlike any other mammal, because their hollow, "air-filled, quill-like hair" acts as a supporting "life jacket". A darker belly color may be caused by two mutations of MC1R. They appear to be more common in domestic reindeer herds. Heat exchange Blood moving into the legs is cooled by blood returning to the body in a countercurrent heat exchange (CCHE), a highly efficient means of minimizing heat loss through the skin's surface. In the CCHE mechanism, in cold weather, blood vessels are closely knotted and intertwined with arteries to the skin and appendages that carry warm blood with veins returning to the body that carry cold blood causing the warm arterial blood to exchange heat with the cold venous blood. In this way, their legs for example are kept cool, maintaining the core body temperature nearly higher with less heat lost to the environment. Heat is thus recycled instead of being dissipated. The "heart does not have to pump blood as rapidly in order to maintain a constant body core temperature and thus, metabolic rate." CCHE is present in animals like reindeer, fox and moose living in extreme conditions of cold or hot weather as a mechanism for retaining the heat in (or out of) the body. These are countercurrent exchange systems with the same fluid, usually blood, in a circuit, used for both directions of flow. Reindeer have specialized counter-current vascular heat exchange in their nasal passages. Temperature gradient along the nasal mucosa is under physiological control. Incoming cold air is warmed by body heat before entering the lungs and water is condensed from the expired air and captured before the reindeer's breath is exhaled, then used to moisten dry incoming air and possibly be absorbed into the blood through the mucous membranes. Like moose, caribou have specialized noses featuring nasal turbinate bones that dramatically increase the surface area within the nostrils. Hooves The reindeer has large feet with crescent-shaped cloven hooves for walking in snow or swamps. According to the Species at Risk Public Registry (SARA), woodland Size The females (or "cows" as they are often called) usually measure in length and weigh . The males (or "bulls" as they are often called) are typically larger (to an extent which varies between the different species and subspecies), measuring in length and usually weighing . The shoulder height is usually , and the tail is long. The reindeer from Svalbard are the smallest of all. They are also relatively short-legged and may have a shoulder height of as little as , thereby following Allen's rule. Clicking sound The knees of many species and subspecies of reindeer are adapted to produce a clicking sound as they walk. The sounds originate in the tendons of the knees and may be audible from several hundred meters away. The frequency of the knee-clicks is one of a range of signals that establish relative positions on a dominance scale among reindeer. "Specifically, loud knee-clicking is discovered to be an honest signal of body size, providing an exceptional example of the potential for non-vocal acoustic communication in mammals." The sound is made when a reindeer is walking or running, occurring when the full weight of the foot is on the ground or just after it is relieved of the weight. It has been proposed that UV flashes on power lines are responsible for reindeer avoiding power lines because "...in darkness these animals see power lines not as dim, passive structures but, rather, as lines of flickering light stretching across the terrain." In 2023, researchers studying reindeer living in Cairngorms National Park, Scotland, suggested that UV visual sensitivity in reindeer helps them detect UV-absorbing lichens against a background of UV-reflecting snows. The tapetum lucidum of Arctic reindeer eyes changes in color from gold in summer to blue in winter to improve their vision during times of continuous darkness, and perhaps enable them to better spot predators. == Biology and behaviors ==
Biology and behaviors
Seasonal body composition Reindeer have developed adaptations for optimal metabolic efficiency during warm months as well as for during cold months. The body composition of reindeer varies highly with the seasons. Of particular interest is the body composition and diet of breeding and non-breeding females between the seasons. Breeding females have more body mass than non-breeding females between the months of March and September with a difference of around more than non-breeding females. From November to December, non-breeding females have more body mass than breeding females, as non-breeding females are able to focus their energies towards storage during colder months rather than lactation and reproduction. Body masses of both breeding and non-breeding females peaks in September. During the months of March through April, breeding females have more fat mass than the non-breeding females with a difference of almost . After this, however, non-breeding females on average have a higher body fat mass than do breeding females. The environmental variations play a large part in reindeer nutrition, as winter nutrition is crucial to adult and neonatal survival rates. Lichens are a staple during the winter months as they are a readily available food source, which reduces the reliance on stored body reserves. In a study of seasonal light-dark cycles on sleep patterns of female reindeer, researchers performed non-invasive electroencephalography (EEG) on reindeer kept in a stable at the UiT The Arctic University of Norway. The EEG recordings showed that: (1) the more time reindeer spend ruminating, the less time they spend in non-rapid eye movement sleep (NREM sleep); and (2) reindeer's brainwaves during rumination resemble the brainwaves present during NREM sleep. These results suggest that, by reducing the time requirement for NREM sleep, reindeer are able to spend more time feeding during the summer months, when food is abundant. Reproduction and life cycle Reindeer mate in late September to early November, and the gestation period is about 228–234 days. During the mating season, bulls battle for access to cows. Two bulls will lock each other's antlers together and try to push each other away. The most dominant bulls can collect as many as 15–20 cows to mate with. A bull will stop eating during this time and lose much of his body fat reserves. To calve, "females travel to isolated, relatively predator-free areas such as islands in lakes, peatlands, lake-shores, or tundra." Dominant bulls, those with larger body size and antler racks, inseminate more than one cow a season. Social structure, migration and range loads. Severely infected individuals are weak and probably have shortened lifespans, but parasite levels vary between populations. Infections create an effect known as culling: infected migrating animals are less likely to complete the migration. Normally travelling about a day while migrating, the caribou can run at speeds of . Young calves can already outrun an Olympic sprinter when only 1 day old. During the spring migration, smaller herds will group together to form larger herds of 50,000 to 500,000 animals, but during autumn migrations, the groups become smaller and the reindeer begin to mate. During winter, reindeer travel to forested areas to forage under the snow. By spring, groups leave their winter grounds to go to the calving grounds. A reindeer can swim easily and quickly, normally at about but, if necessary, at and migrating herds will not hesitate to swim across a large lake or broad river. The barren-ground caribou form large herds and undertake lengthy seasonal migrations from winter feeding grounds in taiga to spring calving grounds and summer range in the tundra. The migrations of the Porcupine herd of barren-ground caribou are among the longest of any mammal. Unlike the individual-tending mating system, aggregated rutting, synchronized calving and aggregated post-calving of barren-ground caribou, Greenland caribou have a harem-defense mating system and dispersed calving and they do not aggregate. Finnish forest reindeer (R. t. fennicus) were formerly distributed in most of the coniferous forest zones south of the tree line, including some mountains, but are now spottily distributed within this zone. As an adaptation to their Arctic environment, they have lost their circadian rhythm. == Ecology ==
Ecology
Distribution and habitat , Finland in Mongolia Originally, the reindeer was found in Scandinavia, Eastern Europe, Greenland, Russia, Mongolia and northern China north of the 50th latitude. In North America, it was found in Canada, Alaska, and the northern contiguous United States from Maine to Washington. In the 19th century, it was still present in southern Idaho. Even in historical times, it probably occurred naturally in Ireland, and it is believed to have lived in Scotland until the 12th century, when the last reindeer were hunted in Orkney. During the Late Pleistocene Epoch, reindeer occurred further south in North America, such as in Nevada, Tennessee, and Alabama, and as far south as Spain in Europe. Though their range retreated northwards during the terminal Pleistocene, reindeer returned to Northern Europe during the Younger Dryas. Today, wild reindeer have disappeared from these areas, especially from the southern parts, where it vanished almost everywhere. Large populations of wild reindeer are still found in Norway, Finland, Siberia, Greenland, Alaska and Canada. According to Grubb (2005), Rangifer is "circumboreal in the tundra and taiga" from "Svalbard, Norway, Finland, Russia, Alaska (USA) and Canada including most Arctic islands, and Greenland, south to northern Mongolia, China (Inner Mongolia), Sakhalin Island, and USA (northern Idaho and Great Lakes region)." Reindeer were introduced to, and are feral in, "Iceland, Kerguelen Islands, South Georgia Island, Pribilof Islands, St. Matthew Island"; There is strong regional variation in Rangifer herd size. There are large population differences among individual herds and the size of individual herds has varied greatly since 1970. The largest of all herds (in Taimyr, Russia) has varied between 400,000 and 1,000,000; the second largest herd (at the George River in Canada) has varied between 28,000 and 385,000. While Rangifer is a widespread and numerous genus in the northern Holarctic, being present in both tundra and taiga (boreal forest), Diet Reindeer are ruminants, having a four-chambered stomach. They mainly eat lichens in winter, especially reindeer lichen (Cladonia rangiferina); they are the only large mammal able to metabolize lichen. Many lichens synthesize toxic phenolic compounds, such as usnic acid, that protect them from UV radiation and herbivory. Reindeer can eat lichen because their rumen contains a reindeer-specific bacterium called Eubacterium rangiferina. This bacterium detoxifies usnic acid and may use it as an energy source. Unlike other bacteria present in the rumen, E. rangiferina can grow in the presence of usnic acid and rapidly degrades usnic acid in reindeer stomachs. Reindeer also eat the leaves of willows and birches, as well as sedges and grasses, especially in the warmer months. Reindeer are osteophagous; they are known to gnaw and partly consume shed antlers as a dietary supplement and in some extreme cases will cannibalise each other's antlers before shedding. There is also some evidence to suggest that on occasion, especially in the spring when they are nutritionally stressed, they will feed on small rodents (such as lemmings), fish (such as the Arctic char (Salvelinus alpinus)), and bird eggs. Reindeer herded by the Chukchis have been known to devour mushrooms enthusiastically in late summer. During the Arctic summer, when there is continuous daylight, reindeer change their sleeping pattern from one synchronised with the sun to an ultradian pattern, in which they sleep when they need to digest food. δ13CC values indicate reindeer living in the region around Biśnik Cave exhibited minimal ecological change during the transition from MIS 3 to MIS 2. Dental mesowear indicates that during the Late Pleistocene, reindeer living in central Alaska had highly abrasive diets similar to wild horses. Predators A variety of predators prey heavily on reindeer, including overhunting by people in some areas, which contributes to the decline of populations. Wolverines will take newborn calves or birthing cows, as well as (less commonly) infirm adults. Brown bears and polar bears prey on reindeer of all ages but, like wolverines, are most likely to attack weaker animals, such as calves and sick reindeer, since healthy adult reindeer can usually outpace a bear. The gray wolf is the most effective natural predator of adult reindeer and sometimes takes large numbers, especially during the winter. Some gray wolf packs, as well as individual grizzly bears in Canada, may follow and live off of a particular reindeer herd year-round. In 2020, scientists on Svalbard witnessed, and were able to film for the first time, a polar bear attack reindeer, driving one into the ocean, where the polar bear caught up with and killed it. The same bear successfully repeated this hunting technique the next day. On Svalbard, reindeer remains account for 27.3% in polar bear scats, suggesting that they "may be a significant part of the polar bear's diet in that area". Additionally, as carrion, reindeer may be scavenged opportunistically by red and Arctic foxes, various species of eagles, hawks and falcons, and common ravens. Bloodsucking insects, such as mosquitoes, black flies, and especially the reindeer warble fly or reindeer botfly (Hypoderma tarandi) and the reindeer nose botfly (Cephenemyia trompe), are a plague to reindeer during the summer and can cause enough stress to inhibit feeding and calving behaviors. An adult reindeer will lose perhaps about of blood to biting insects for every week it spends in the tundra. Reindeer are good swimmers and, in one case, the entire body of a reindeer was found in the stomach of a Greenland shark (Somniosus microcephalus), a species found in the far North Atlantic. Other threats White-tailed deer (Odocoileus virginianus) commonly carry meningeal worm or brainworm (Parelaphostrongylus tenuis), a nematode parasite that causes reindeer, moose (Alces alces), elk (Cervus canadensis), and mule deer (Odocoileus hemionus) to develop fatal neurological symptoms which include a loss of fear of humans. White-tailed deer that carry this worm are partially immune to it. == Conservation ==
Conservation
Current status According to the IUCN, Rangifer tarandus, as a species, is not endangered because of its overall large population and its widespread range, but, as of 2015, the IUCN has classified the reindeer as Vulnerable due to an observed population decline of 40% over the last +25 years. and the East Greenland caribou both became extinct in the early 20th century, the Peary caribou is designated as Endangered, the boreal woodland caribou is designated as Threatened and some individual populations are endangered as well. While the barren-ground caribou is not designated as Threatened, many individual herds — including some of the largest — are declining and there is much concern at the local level. Grant's caribou, a small, pale subspecies endemic to the western end of the Alaska Peninsula and the adjacent islands, In NWT, Dolphin-Union caribou were listed as Special Concern under the NWT Species at Risk (NWT) Act (2013). Both the Selkirk Mountains caribou (Southern Mountain population DU9) and the Rocky Mountain caribou (Central Mountain population DU8) are classified as Endangered in Canada in regions such as southeastern British Columbia at the Canada–United States border, along the Columbia and Kootenay Rivers and around Kootenay Lake. Rocky Mountain caribou are extirpated from Banff National Park, but a small population remains in Jasper National Park and in mountain ranges to the northwest into British Columbia. Montane caribou are now considered extirpated in the contiguous United States, including Washington and Idaho. Osborn's caribou (Northern Mountain population DU7) is classified as Threatened in Canada. In Eurasia, the Sakhalin reindeer is extinct (and has been replaced by domestic reindeer) and reindeer on most of the Novaya Zemlya islands have also been replaced by domestic reindeer, although some wild reindeer still persist on the northern islands. Boreal woodland caribou were designated as Threatened in 2002 by the Committee on the Status of Endangered Wildlife in Canada, (COSEWIC). Environment Canada reported in 2011 that there were approximately 34 000 boreal woodland caribou in 51 ranges remaining in Canada (Environment Canada, 2011b). "According to Geist, the "woodland caribou is highly endangered throughout its distribution right into Ontario." In 2002, the Atlantic-Gaspésie population DU11 of the boreal woodland caribou was designated as Endangered by COSEWIC. The small isolated population of 200 animals was at risk from predation and habitat loss. Peary caribou In 1991, COSEWIC assigned "endangered status" to the Banks Island and High Arctic populations of the Peary caribou. The Low Arctic population of the Peary caribou was designated as Threatened. In 2004, all three were designated as "endangered." In 2015, COSEWIC returned the status to Threatened. == Relationship with humans ==
Relationship with humans
Arctic peoples have depended on caribou for food, clothing, and shelter. European prehistoric cave paintings represent both tundra and forest forms, the latter either the Finnish forest reindeer or the narrow-nosed reindeer, an eastern Siberia forest form. Reindeer have been domesticated at least two and probably three times, in each case from wild Eurasian tundra reindeer after the Last Glacial Maximum (LGM). the Tuvans, Todzhans, Tofa (Tofalars in the Irkutsk Region), the Soyots (the Republic of Buryatia), and the Dukha (also known as Tsaatan, the Khubsugul) in the Province of Mongolia. The Sámi (Sápmi) have also depended on reindeer herding and fishing for centuries. In Sápmi, reindeer are used to pull a pulk, a Nordic sled. Furthermore, Lapin Poron liha, fresh reindeer meat completely produced and packed in Finnish Lapland, is protected in Europe with PDO classification. Reindeer antlers are powdered and sold as an aphrodisiac, or as a nutritional or medicinal supplement, to Asian markets. The blood of the caribou was supposedly mixed with alcohol as drink by hunters and loggers in colonial Quebec to counter the cold. This drink is now enjoyed without the blood as a wine and whiskey drink known as Caribou. Indigenous North Americans Caribou are still hunted in Greenland and in North America. In the traditional lifestyles of some of Canada's Inuit peoples and northern First Nations peoples, Alaska Natives, and the Kalaallit of Greenland, caribou is an important source of food, clothing, shelter and tools. made of caribou skin The Caribou Inuit are inland-dwelling Inuit in present-day Nunavut's Kivalliq Region (formerly the Keewatin Region), Canada. They subsisted on caribou year-round, eating dried caribou meat in the winter. The Ahiarmiut are Caribou Inuit that followed the Qamanirjuaq barren-ground caribou herd. There is an Inuit saying in the Kivalliq Region: The Gwichʼin, an indigenous people of northwestern Canada and northeastern Alaska, have been dependent on the international migratory Porcupine caribou herd for millennia. To them, caribou — vadzaih — is the cultural symbol and a keystone subsistence species of the Gwich'in, just as the American buffalo is to the Plains Native Americans. Innovative language revitalisation projects are underway to document the language and to enhance the writing and translation skills of younger Gwich'in speakers. In one project, lead research associate and fluent speaker Gwich'in elder Kenneth Frank works with linguists who include young Gwich'in speakers affiliated with the Alaska Native Language Center at the University of Alaska in Fairbanks to document traditional knowledge of caribou anatomy. The main goal of the research was to "elicit not only what the Gwich'in know about caribou anatomy, but how they see caribou and what they say and believe about caribou that defines themselves, their dietary and nutritional needs, and their subsistence way of life." Indigenous Eurasians Reindeer herding has been vital for the subsistence of several Eurasian nomadic indigenous peoples living in the circumpolar Arctic zone such as the Sámi, Nenets, and Komi. Reindeer are used to provide renewable sources and reliable transportation. In Mongolia, the Dukha are known as the reindeer people. They are credited as one of the world's earliest domesticators. The Dukha diet consists mainly of reindeer dairy products. Reindeer husbandry is common in northern Fennoscandia (northern Norway, Sweden and Finland) and the Russian North. In some human groups such as the Eveny, wild reindeer and domestic reindeer are treated as different kinds of beings. Husbandry , Russia, late 19th-century photochrom , Norway, 19th century The reindeer is the only successfully semi-domesticated deer on a large scale in the world. Reindeer in northern Fennoscandia (northern Norway, Sweden and Finland) as well in the Kola Peninsula and Yakutia in Russia, are mostly semi-domesticated reindeer, ear-marked by their owners. Some reindeer in the area are truly domesticated, mostly used as draught animals (nowadays commonly for tourist entertainment and races, traditionally important for the nomadic Sámi). Domestic reindeer have also been used for milk, e.g., in Norway. There are only two genetically pure populations of wild reindeer in Northern Europe: wild mountain reindeer (R. t. tarandus) that live in central Norway, with a population in 2007 of between 6,000 and 8,400 animals; and wild Finnish forest reindeer (R. t. fennicus) that live in central and eastern Finland and in Russian Karelia, with a population of about 4,350, plus 1,500 in Arkhangelsk Oblast and 2,500 in Komi. East of Arkhangelsk, both wild Siberian tundra reindeer (R. t. sibiricus) (some herds are very large) and domestic reindeer (R. t. domesticus) occur with almost no interbreeding by wild reindeer into domestic clades and none the other way (Kharzinova et al. 2018; Rozhkov et al. 2020 Reindeer have been herded for centuries by several Arctic and sub-Arctic peoples, including the Sámi, the Nenets and the Yakuts. They are raised for their meat, hides and antlers and, to a lesser extent, for milk and transportation. Reindeer are not considered fully domesticated, as they generally roam free on pasture grounds. In traditional nomadic herding, reindeer herders migrate with their herds between coastal and inland areas according to an annual migration route and herds are keenly tended. However, reindeer were not bred in captivity, though they were tamed for milking as well as for use as draught animals or beasts of burden. Millais (1915), The use of reindeer for transportation is common among the nomadic peoples of the Russian North (but not anymore in Scandinavia). Although a sled drawn by 20 reindeer will cover no more than a day (compared to on foot, by a dog sled loaded with cargo and by a dog sled without cargo), it has the advantage that the reindeer will discover their own food, while a pack of 5–7 sled dogs requires of fresh fish a day. The use of reindeer as semi-domesticated livestock in Alaska was introduced in the late 19th century by the United States Revenue Cutter Service, with assistance from Sheldon Jackson, as a means of providing a livelihood for Alaska Natives. Reindeer were imported first from Siberia and later also from Norway. A regular mail run in Wales, Alaska, used a sleigh drawn by reindeer. In Alaska, reindeer herders use satellite telemetry to track their herds, using online maps and databases to chart the herd's progress. Domestic reindeer are mostly found in northern Fennoscandia and the Russian North, with a herd of approximately 150–170 reindeer living around the Cairngorms region in Scotland. The last remaining wild tundra reindeer in Europe are found in portions of southern Norway. The International Centre for Reindeer Husbandry (ICR), a circumpolar organisation, was established in 2005 by the Norwegian government. ICR represents over 20 indigenous reindeer peoples and about 100,000 reindeer herders in nine different national states. In Finland, there are about 6,000 reindeer herders, most of whom keep small herds of less than 50 reindeer to raise additional income. With 185,000 reindeer (), the industry produces of reindeer meat and generates 35 million euros annually. 70% of the meat is sold to slaughterhouses. Reindeer herders are eligible for national and EU agricultural subsidies, which constituted 15% of their income. Reindeer herding is of central importance for the local economies of small communities in sparsely populated rural Sápmi. Currently, many reindeer herders are heavily dependent on diesel fuel to provide for electric generators and snowmobile transportation, although solar photovoltaic systems can be used to reduce diesel dependency. File:Carta Marina - milking reindeer.jpg|Milking File:Carta Marina - reindeer crossing a frozen lake.jpg|Crossing frozen water File:Carta Marina - reindeer-drawn waggon with bowman.jpg|Drawing a wagon File:Carta Marina - reindeer-drawn sled.jpg|Drawing a one-man sled File:Carta Marina - reindeer-mounted warriors.jpg|Reindeer-mounted cavalry History Reindeer hunting by humans has a very long history. Both Aristotle and Theophrastus have short accounts – probably based on the same source – of an ox-sized deer species, named tarandos, living in the land of the Bodines in Scythia, which was able to change the colour of its fur to obtain camouflage. The latter is probably a misunderstanding of the seasonal change in reindeer fur colour. The descriptions have been interpreted as being of reindeer living in the southern Ural Mountains in c. 350 BC. A deer-like animal described by Julius Caesar in his Commentarii de Bello Gallico (chapter 6.26) from the Hercynian Forest in the year 53 BC is most certainly to be interpreted as a reindeer: According to Olaus Magnus's Historia de Gentibus Septentrionalibus – printed in Rome in the year 1555 – Gustav I of Sweden sent 10 reindeer to Albert, Duke of Prussia, in the year 1533. It may be these animals that Conrad Gessner had seen or heard of. During World War II, the Soviet Army used reindeer as pack animals to transport food, ammunition and post from Murmansk to the Karelian front and bring wounded soldiers, pilots and equipment back to the base. About 6,000 reindeer and more than 1,000 reindeer herders were part of the operation. Most herders were Nenets, who were mobilised from the Nenets Autonomous Okrug, but reindeer herders from the Murmansk, Arkhangelsk and Komi regions also participated. In the FIS Alpine Ski World Cup event held in Levi, Finland each year, the winner of the women's slalom event is awarded a reindeer. The prize is largely symbolic, as all the reindeer awarded continue living in on a farm in Finland. Santa Claus Around the world, public interest in reindeer peaks during the Christmas season. According to Western folklore, Santa Claus's sleigh is pulled by flying reindeer. These reindeer were first named in the 1823 poem "A Visit from St. Nicholas", though the story originates earlier. Mythology and art Among the Inuit, there is a story of the origin of the caribou: Inuit artists from the Barrenlands incorporate depictions of caribou—and items made from caribou antlers and skin—in carvings, drawings, prints and sculpture. Contemporary Canadian artist Brian Jungen, of Dane-zaa First Nations ancestry, commissioned an installation entitled "The ghosts on top of my head" (2010–11) in Banff, Alberta, which depicts the antlers of caribou, elk and moose. Tomson Highway, CM is a Canadian and Cree playwright, novelist, and children's author, who was born in a remote area north of Brochet, Manitoba. The Canadian 25-cent coin or "quarter" features a depiction of a caribou on one face. The caribou is the official provincial animal of Newfoundland and Labrador, Canada, and appears on the coat of arms of Nunavut. A caribou statue was erected at the centre of the Beaumont-Hamel Newfoundland Memorial, marking the spot in France where hundreds of soldiers from Newfoundland were killed and wounded in World War I. There is a replica in Bowring Park in St. John's, Newfoundland's capital city. Two municipalities in Finland have reindeer motifs in their coats-of-arms: Kuusamo has a running reindeer; and Inari has a fish with reindeer antlers. File:Kuusamo.vaakuna.svg|Coat of arms of Kuusamo|alt=Coat of arms of Kuusamo features a male File:Inari.vaakuna.svg|Coat of arms of Inari|alt=Coat of arms of Inari, a fish with antlers == See also ==
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