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The reindeer or caribou (''Rangifer tarandus'') is a species of
deer A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) ...
with
circumpolar distribution A circumpolar distribution is any range (biology), range of a taxon that occurs over a wide range of longitudes but only at high latitudes; such a range therefore extends all the way around either the North Pole or the South Pole. Taxa that are al ...
, native to
Arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
,
subarctic The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of hemiboreal regions and covering much of Alaska, Canada, Iceland, the north of Fennoscandia, Northwestern Russia, Siberia, and the Cair ...
,
tundra In physical geography, a tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. There are three regions and associated types of tundra: #Arctic, Arctic, Alpine tundra, Alpine, and #Antarctic ...
,
boreal Boreal, northern, of the north. Derived from the name of the god of the north wind from Ancient Greek civilisation, Boreas (god), Boreas. It may also refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernand ...
, 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 into six distinct species over their range. Reindeer occur in both migratory and
sedentary Sedentary lifestyle is a lifestyle type, in which one is physically inactive and does little or no physical movement and/or exercise. A person living a sedentary lifestyle is often sitting or lying down while engaged in an activity like soc ...
populations, and their herd sizes vary greatly in different regions. The tundra subspecies are adapted for extreme cold, and some are adapted for long-distance migration. Reindeer vary greatly in size and color from the smallest, the
Svalbard reindeer The Svalbard reindeer (''Rangifer tarandus platyrhynchus'') is a small subspecies or species of reindeer found on the Svalbard archipelago of Norway. Males average in weight, females , while for other reindeer generally body mass is for males a ...
(''R.'' (''t.'') ''platyrhynchus''), to the largest, Osborn's caribou (''R. t. osborni''). Although reindeer are quite numerous, some species and subspecies are in decline and considered vulnerable. They are unique among deer (Cervidae) in that females may have
antler Antlers are extensions of an animal's skull found in members of the Cervidae (deer) Family (biology), family. Antlers are a single structure composed of bone, cartilage, fibrous tissue, skin, nerves, and blood vessels. They are generally fo ...
s, although the prevalence of antlered females varies by subspecies. Reindeer are the only successfully semi-domesticated deer on a large scale in the world. Both wild and domestic reindeer have been an important source of food, clothing, and shelter for Arctic people from prehistorical times. They are still herded and hunted today. In some traditional
Christmas Christmas is an annual festival commemorating Nativity of Jesus, the birth of Jesus Christ, observed primarily on December 25 as a Religion, religious and Culture, cultural celebration among billions of people Observance of Christmas by coun ...
legends,
Santa Claus's reindeer In traditional Western festive legend and popular culture, Santa Claus's reindeer are said to pull a sleigh through the night sky to help Santa Claus deliver gifts to children on Christmas Eve. While various legends offer differing details ...
pull a sleigh through the night sky to help Santa Claus deliver gifts to good children on Christmas Eve.


Description

Names follow international conventionMattioli, S. (2011). "Caribou (''Rangifer tarandus'')", pp. 431–432 in: ''Handbook of the Mammals of the World Vol. 2: Hoofed Mammals''. Lynx Edicions, Barcelona. before the recent revision (see below). Reindeer / caribou (''Rangifer'') vary in size from the smallest, the
Svalbard reindeer The Svalbard reindeer (''Rangifer tarandus platyrhynchus'') is a small subspecies or species of reindeer found on the Svalbard archipelago of Norway. Males average in weight, females , while for other reindeer generally body mass is for males a ...
(''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 Yukon () is a Provinces and territories of Canada, territory of Canada, bordering British Columbia to the south, the Northwest Territories to the east, the Beaufort Sea to the north, and the U.S. state of Alaska to the west. It is Canada’s we ...
, the
Northwest Territories The Northwest Territories is a federal Provinces and territories of Canada, territory of Canada. At a land area of approximately and a 2021 census population of 41,070, it is the second-largest and the most populous of Provinces and territorie ...
and
Nunavut Nunavut is the largest and northernmost Provinces and territories of Canada#Territories, territory of Canada. It was separated officially from the Northwest Territories on April 1, 1999, via the ''Nunavut Act'' and the Nunavut Land Claims Agr ...
throughout the
tundra In physical geography, a tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. There are three regions and associated types of tundra: #Arctic, Arctic, Alpine tundra, Alpine, and #Antarctic ...
,
taiga Taiga or tayga ( ; , ), also known as boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruces, and larches. The taiga, or boreal forest, is the world's largest land biome. In North A ...
(boreal forest) and south through the Canadian
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in great-circle distance, straight-line distance from the northernmost part of Western Can ...
. Of the eight subspecies classified by Harding (2022) into the Arctic caribou (''R. arcticus''), the migratory mainland
barren-ground caribou The barren-ground caribou (''Rangifer tarandus groenlandicus''; but subject to a recent taxonomic revision) is a subspecies of the reindeer (or the caribou in North America) that is found in the Canadian territories of Nunavut and the Northwest ...
of
Arctic Alaska Arctic Alaska or Far North Alaska is a region of the United States, U.S. U.S. state, state of Alaska generally referring to the northern areas on or close to the Arctic Ocean. It commonly includes North Slope Borough, Alaska, North Slope Boro ...
and
Northern Canada Northern Canada (), colloquially the North or the Territories, is the vast northernmost region of Canada, variously defined by geography and politics. Politically, the term refers to the three Provinces_and_territories_of_Canada#Territories, terr ...
(''R. t. arcticus''), summer in tundra and winter in taiga, a transitional forest zone between boreal forest and tundra; the nomadic
Peary caribou The Peary caribou (''Rangifer arcticus pearyi'') is a subspecies of caribou found in the Canadian high Arctic islands of Nunavut and the Northwest Territories in Canada. They are the smallest of the North American caribou, with the females wei ...
(''R. t. pearyi'') lives in the
polar desert Polar deserts are the regions of Earth that fall under an ice cap climate (''EF'' under the Köppen classification). Despite rainfall totals low enough to normally classify as a desert, polar deserts are distinguished from true deserts (' or ' un ...
of the high
Arctic Archipelago The Arctic Archipelago, also known as the Canadian Arctic Archipelago, is an archipelago lying to the north of the Canadian continental mainland, excluding Greenland (an autonomous territory of the Danish Realm, which is, by itself, much larger ...
and Grant's caribou (''R. t. granti'' also called the
Porcupine caribou The Porcupine caribou is a herd or ecotype of the mainland barren-ground caribou (''Rangifer arcticus arcticus'', Synonym (taxonomy), syn. ''R. tarandus groenlandicus''Harding LE (2022) Available names for Rangifer (Mammalia, Artiodactyla, Cervid ...
) lives in the western end of the
Alaska Peninsula The Alaska Peninsula (also called Aleut Peninsula or Aleutian Peninsula, ; Sugpiaq language, Sugpiaq: ''Aluuwiq'', ''Al'uwiq'') is a peninsula extending about to the southwest from the mainland of Alaska and ending in the Aleutian Islands. T ...
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 Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
. The extinct insular
Queen Charlotte Islands caribou The Dawson's caribou, also known as the Queen Charlotte Islands caribou (''Rangifer tarandus dawsoni'') was a population of woodland caribou that once lived on Graham Island, the largest of the islands within the Haida Gwaii archipelago, locate ...
(''R. t. dawsoni''), lived on
Graham Island Graham Island () is the largest island in the Haida Gwaii archipelago (previously known as the Queen Charlotte Islands), lying off the mainland coast of British Columbia, Canada. It is separated by the narrow Skidegate Channel from the other pr ...
in
Haida Gwaii Haida Gwaii (; / , literally "Islands of the Haida people"), previously known as the Queen Charlotte Islands, is an archipelago located between off the British Columbia Coast, northern Pacific coast in the Canadian province of British Columbia ...
(formerly known as the Queen Charlotte Islands). The
boreal woodland caribou The boreal woodland caribou (''Rangifer tarandus caribou''; but subject to a recent taxonomic revision. See Reindeer: Taxonomy), also known as Eastern woodland caribou, boreal forest caribou and forest-dwelling caribou, is a North American subsp ...
(''R. t. caribou''), lives in the boreal forest of northeastern Canada: the Labrador or Ungava caribou of
northern Quebec Northern Quebec () is a geographic term denoting the northerly, more remote and less populated parts of the Canada, Canadian province of Quebec.Alexandre Robaey"Charity group works with Indigenous communities to feed Northern Quebec's 'wandering dog ...
and northern
Labrador Labrador () is a geographic and cultural region within the Canadian province of Newfoundland and Labrador. It is the primarily continental portion of the province and constitutes 71% of the province's area but is home to only 6% of its populatio ...
(''R. t. caboti''), and the Newfoundland caribou of
Newfoundland Newfoundland and Labrador is the easternmost province of Canada, in the country's Atlantic region. The province comprises the island of Newfoundland and the continental region of Labrador, having a total size of . As of 2025 the population ...
(''R. t. terranovae'') have been found to be genetically in the woodland caribou lineage. In Eurasia, both wild and domestic reindeer are distributed across the tundra and into the taiga. Eurasian mountain reindeer (''R. t. tarandus'') are close to North American caribou genetically and visually, but with sufficient differences to warrant division into two species. The unique, insular Svalbard reindeer inhabits the
Svalbard Archipelago Svalbard ( , ), previously known as Spitsbergen or Spitzbergen, is a Norway, Norwegian archipelago that lies at the convergence of the Arctic Ocean with the Atlantic Ocean. North of continental Europe, mainland Europe, it lies about midway be ...
. The
Finnish forest reindeer The Finnish forest reindeer ''(Rangifer tarandus fennicus'' (Finnish:'' metsäpeura'', Russian: ''лесной северный олень''), also known as Eurasian or European forest reindeer is a rare subspecies of the reindeer native to Finla ...
(''R. t. fennicus'') is spottily distributed in the coniferous forest zones from Finland to east of
Lake Baikal Lake Baikal is a rift lake and the deepest lake in the world. It is situated in southern Siberia, Russia between the Federal subjects of Russia, federal subjects of Irkutsk Oblast, Irkutsk Oblasts of Russia, Oblast to the northwest and the Repu ...
: the Siberian forest reindeer (''R. t. valentinae'', formerly called the Busk Mountains reindeer (''R. t. buskensis'') by American taxonomists) occupies the Altai and
Ural Mountains The Ural Mountains ( ),; , ; , or simply the Urals, are a mountain range in Eurasia that runs north–south mostly through Russia, from the coast of the Arctic Ocean to the river Ural (river), Ural and northwestern Kazakhstan.
. Male ("bull") and female ("cow") reindeer can grow antlers annually, although the proportion of females that grow antlers varies greatly between populations. Antlers are typically larger on males. Antler architecture varies by species and subspecies and, together with pelage differences, can often be used to distinguish between species and subspecies (see illustrations in Geist, 1991 and Geist, 1998).


Status

About 25,000 mountain reindeer (''R. t. tarandus'') still live in the mountains of Norway, notably in
Hardangervidda Hardangervidda () is a mountain plateau ( Norwegian: ''vidde'') in central southern Norway, covering parts of Vestland, Telemark, and Buskerud counties. It is the largest plateau of its kind in Europe, with a cold year-round alpine climate, and o ...
. In Sweden there are approximately 250,000 reindeer in herds managed by
Sámi Acronyms * SAMI, ''Synchronized Accessible Media Interchange'', a closed-captioning format developed by Microsoft * Saudi Arabian Military Industries, a government-owned defence company * South African Malaria Initiative, a virtual expertise ...
villages. Russia manages 19 herds of Siberian tundra reindeer (''R. t. sibiricus'') that total about 940,000.Mizin, I.A. (2018) The current state of the wild reindeer in Russia - general overview of the situation. Barents office of WWF Russia Russian Arctic National Park for World Wildlife Fund, Arkhangelsk, Russia, 8 pp. 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 The Sea of Okhotsk; Historically also known as , or as ; ) is a marginal sea of the western Pacific Ocean. It is located between Russia's Kamchatka Peninsula on the east, the Kuril Islands on the southeast, Japan's island of Hokkaido on the sou ...
which, however, are indistinguishable genetically from the Jano-Indigirka,
East Siberian taiga The East Siberian taiga ecoregion, in the taiga and boreal forests biome, is a very large biogeographic region in eastern Russia. Setting This vast ecoregion is located in the heart of Siberia, stretching over 20° of latitude and 50° of longit ...
and Chukotka populations of ''R. t. sibiricus''. Siberian tundra reindeer herds have been in decline but are stable or increasing since 2000. Insular (island) reindeer, classified as the Novaya Zemlya reindeer (''R. t. pearsoni'') occupy several island groups: the
Novaya Zemlya Novaya Zemlya (, also , ; , ; ), also spelled , is an archipelago in northern Russia. It is situated in the Arctic Ocean, in the extreme northeast of Europe, with Cape Flissingsky, on the northern island, considered the extreme points of Europe ...
Archipelago (about 5,000 animals at last count, but most of these are either domestic reindeer or domestic-wild hybrids), the
New Siberia New Siberia or Novaya Sibir (; , , ; ) is the easternmost of the Anzhu Islands, the northern subgroup of the New Siberian Islands lying between the Laptev Sea and East Siberian Sea. Its area of approximately places it the 102nd largest islan ...
Archipelago (about 10,000 to 15,000), and
Wrangel Island Wrangel Island (, ; , , ) is an island of the Chukotka Autonomous Okrug, Russia. It is the List of islands by area, 92nd-largest island in the world and roughly the size of Crete. Located in the Arctic Ocean between the Chukchi Sea and East Si ...
(200 to 300 feral domestic reindeer). What was once the second largest herd is the migratory Labrador caribou (''R. t. caboti'')
George River herd George may refer to: Names * George (given name) * George (surname) People * George (singer), American-Canadian singer George Nozuka, known by the mononym George * George Papagheorghe, also known as Jorge / GEØRGE * George, stage name of Giorg ...
in Canada, with former variations between 28,000 and 385,000. As of January 2018, there are fewer than 9,000 animals estimated to be left in the George River herd, as reported by the
Canadian Broadcasting Corporation The Canadian Broadcasting Corporation (), branded as CBC/Radio-Canada, is the Canadian Public broadcasting, public broadcaster for both radio and television. It is a Crown corporation that serves as the national public broadcaster, with its E ...
. ''
The New York Times ''The New York Times'' (''NYT'') is an American daily newspaper based in New York City. ''The New York Times'' covers domestic, national, and international news, and publishes opinion pieces, investigative reports, and reviews. As one of ...
'' reported in April 2018 of the disappearance of the only herd of southern mountain woodland caribou in the
contiguous United States The contiguous United States, also known as the U.S. mainland, officially referred to as the conterminous United States, consists of the 48 adjoining U.S. states and the District of Columbia of the United States in central North America. The te ...
, 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 contiguous United States, also known as the U.S. mainland, officially referred to as the conterminous United States, consists of the 48 adjoining U.S. states and the District of Columbia of the United States in central North America. The te ...
. The
Committee on the Status of Endangered Wildlife in Canada The Committee on the Status of Endangered Wildlife in Canada (COSEWIC, French: Comité sur la situation des espèces en péril au Canada, COSEPAC) is an independent committee of wildlife experts and scientists whose "raison d'être is to identify s ...
(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 The Dawson's caribou, also known as the Queen Charlotte Islands caribou (''Rangifer tarandus dawsoni'') was a population of woodland caribou that once lived on Graham Island, the largest of the islands within the Haida Gwaii archipelago, locate ...
(''R. t. dawsoni'') from western Canada, the Sakhalin reindeer (''R. t. setoni'') from
Sakhalin Sakhalin ( rus, Сахали́н, p=səxɐˈlʲin) is an island in Northeast Asia. Its north coast lies off the southeastern coast of Khabarovsk Krai in Russia, while its southern tip lies north of the Japanese island of Hokkaido. An islan ...
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 The contiguous United States, also known as the U.S. mainland, officially referred to as the conterminous United States, consists of the 48 adjoining U.S. states and the District of Columbia of the United States in central North America. The te ...
from
Maine Maine ( ) is a U.S. state, state in the New England region of the United States, and the northeasternmost state in the Contiguous United States. It borders New Hampshire to the west, the Gulf of Maine to the southeast, and the Provinces and ...
to
Washington Washington most commonly refers to: * George Washington (1732–1799), the first president of the United States * Washington (state), a state in the Pacific Northwest of the United States * Washington, D.C., the capital of the United States ** A ...
. 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 The Committee on the Status of Endangered Wildlife in Canada (COSEWIC, French: Comité sur la situation des espèces en péril au Canada, COSEPAC) is an independent committee of wildlife experts and scientists whose "raison d'être is to identify s ...
(COSEWIC).
Environment and Climate Change Canada Environment and Climate Change Canada (ECCC; )Environment and Climate Change Canada is the applied title under the Federal Identity Program; the legal title is Department of the Environment (). is the department of the Government of Canada res ...
reported in 2011 that there were approximately 34,000
boreal woodland caribou The boreal woodland caribou (''Rangifer tarandus caribou''; but subject to a recent taxonomic revision. See Reindeer: Taxonomy), also known as Eastern woodland caribou, boreal forest caribou and forest-dwelling caribou, is a North American subsp ...
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 The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the stat ...
(IUCN).


Naming

Charles Hamilton Smith Lieutenant-Colonel Charles Hamilton Smith, Royal Guelphic Order, KH, Military Order of William, KW, Royal Society, FRS, Linnean Society of London, FLS (26 December 1776 – 21 September 1859) was a British Army officer, artist, naturalist, ant ...
is credited with the name ''Rangifer'' for the reindeer genus, which
Albertus Magnus Albertus Magnus ( 1200 – 15 November 1280), also known as Saint Albert the Great, Albert of Swabia, Albert von Bollstadt, or Albert of Cologne, was a German Dominican friar, philosopher, scientist, and bishop, considered one of the great ...
used in his , fol. Liber 22, Cap. 268: "Dicitur Rangyfer quasi ramifer". This word may go back to the
Sámi Acronyms * SAMI, ''Synchronized Accessible Media Interchange'', a closed-captioning format developed by Microsoft * Saudi Arabian Military Industries, a government-owned defence company * South African Malaria Initiative, a virtual expertise ...
word .
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
chose the word ''tarandus'' as the specific epithet, making reference to
Ulisse Aldrovandi Ulisse Aldrovandi (11 September 1522 – 4 May 1605) was an Italian naturalist, the moving force behind Bologna's botanical garden, one of the first in Europe. Carl Linnaeus and the comte de Buffon reckoned him the father of natural history stud ...
's fol. 859–863, Cap. 30: De Tarando (1621). However, Aldrovandi and
Conrad Gessner Conrad Gessner (; ; 26 March 1516 – 13 December 1565) was a Swiss physician, naturalist, bibliographer, and philologist. Born into a poor family in Zürich, Switzerland, his father and teachers quickly realised his talents and supported him t ...
thought that ''rangifer'' and ''tarandus'' were two separate animals. In any case, the ''tarandos'' name goes back to
Aristotle Aristotle (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...
and
Theophrastus Theophrastus (; ; c. 371 – c. 287 BC) was an ancient Greek Philosophy, philosopher and Natural history, naturalist. A native of Eresos in Lesbos, he was Aristotle's close colleague and successor as head of the Lyceum (classical), Lyceum, the ...
. The use of the terms ''reindeer'' and ''caribou'' for essentially the same animal can cause confusion, but the ICUN clearly delineates the issue: "Reindeer is the European name for the species of ''Rangifer,'' while in North America, ''Rangifer'' species are known as Caribou." The word ''
reindeer The reindeer or caribou (''Rangifer tarandus'') is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, taiga, boreal, and mountainous regions of Northern Europe, Siberia, and North America. It is the only re ...
'' is an anglicized version of the
Old Norse Old Norse, also referred to as Old Nordic or Old Scandinavian, was a stage of development of North Germanic languages, North Germanic dialects before their final divergence into separate Nordic languages. Old Norse was spoken by inhabitants ...
words ("reindeer") and ("animal") and has nothing to do with reins. The word ''caribou'' comes through French, from the
Mi'kmaq The Mi'kmaq (also ''Mi'gmaq'', ''Lnu'', ''Mi'kmaw'' or ''Mi'gmaw''; ; , and formerly Micmac) are an Indigenous group of people of the Northeastern Woodlands, native to the areas of Canada's Atlantic Provinces, primarily Nova Scotia, New Bru ...
, 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 The Inuvialuit (sing. Inuvialuk; ''the real people'') or Western Canadian Inuit are Inuit who live in the western Canadian Arctic region. They, like all other Inuit, are descendants of the Thule who migrated eastward from Alaska. Their homelan ...
of the western
Canadian Arctic Northern Canada (), colloquially the North or the Territories, is the vast northernmost region of Canada, variously defined by geography and politics. Politically, the term refers to the three territories of Canada: Yukon, Northwest Territories a ...
and
Inuit Inuit (singular: Inuk) are a group of culturally and historically similar Indigenous peoples traditionally inhabiting the Arctic and Subarctic regions of North America and Russia, including Greenland, Labrador, Quebec, Nunavut, the Northwe ...
of the eastern Canadian Arctic, who speak different dialects of the
Inuit languages The Inuit languages are a closely related group of Indigenous languages of the Americas, indigenous American languages traditionally spoken across the North American Arctic and the adjacent subarctic regions as far south as Labrador. The Inuit ...
, both call the barren-ground caribou . The Wekʼèezhìi (
Tłı̨chǫ The Tłı̨chǫ (, ) people, sometimes spelled Tlicho and also known as the Dogrib, are a Dene First Nations people of the Athabaskan-speaking ethnolinguistic group living in the Northwest Territories of Canada. Name The name ''Dogrib' ...
) people, a
Dene The Dene people () are an Indigenous group of First Nations who inhabit the northern boreal, subarctic and Arctic regions of Canada. The Dene speak Northern Athabaskan languages and it is the common Athabaskan word for "people". The term ...
(Athapascan) group, call the Arctic caribou and the boreal woodland caribou . The
Gwichʼin The Gwichʼin (or Kutchin or Loucheux) are an Athabaskan languages, Athabaskan-speaking First Nations in Canada, First Nations people of Canada and an Alaskan Athabaskans, Alaska Native people. They live in the northwestern part of North America ...
(also a Dene group) have over 24 distinct caribou-related words. Reindeer are also called by the
Greenlandic Inuit The Greenlandic Inuit or sometimes simply the Greenlandic are an ethnic group and nation Indigenous peoples of the Americas, indigenous to Greenland, where they constitute the largest ethnic population. They share a common #History, ancestry, ...
and , sometimes , by the
Icelanders Icelanders () are an ethnic group and nation who are native to the island country of Iceland. They speak Icelandic, a North Germanic language. Icelanders established the country of Iceland in mid 930  CE when the (parliament) met for th ...
.


Evolution

The "glacial-interglacial cycles of the
upper Pleistocene The Late Pleistocene is an unofficial age in the international geologic timescale in chronostratigraphy, also known as the Upper Pleistocene from a stratigraphic perspective. It is intended to be the fourth division of the Pleistocene Epoch with ...
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 Last Glacial Maximum refugia were places ('' refugia'') in which humans and other species survived during the Last Glacial Period, around 25,000 to 18,000 years ago. Glacial refugia are areas that climate changes were not as severe, and where sp ...
during the last glacial – the
Wisconsin glaciation The Wisconsin glaciation, also called the Wisconsin glacial episode, was the most recent glacial period of the North American ice sheet complex, peaking more than 20,000 years ago. This advance included the Cordilleran Ice Sheet, which nucleated ...
in North America and the Weichselian glaciation in Eurasia – shaped "intraspecific
genetic variability Genetic variability is either the presence of, or the generation of, genetic differences. It is defined as "the formation of individuals differing in genotype, or the presence of genotypically different individuals, in contrast to environmentally ...
" particularly between the North American and Eurasian parts of the
Arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
. Reindeer / caribou (''Rangifer'') are in the subfamily Odocoileinae, along with roe deer (''
Capreolus ''Capreolus'' is a genus of deer, the roe deer. Etymology English ''roe'' is from Old English ''ra'' or ''rá'', from ''raha'', from Proto-Germanic , cognate to Old Norse ''ra'', Old Saxon ''reho'', Middle Dutch and Dutch ''ree'', Old High ...
''), Eurasian elk / moose (''Alces''), and
water deer The water deer (''Hydropotes inermis'') is a small deer species native to Korea and China. Its prominent tusks, similar to those of musk deer, have led to both subspecies being colloquially named vampire deer in English-speaking areas to which t ...
(''Hydropotes''). These antlered cervids split from the horned ruminants ''
Bos ''Bos'' (from Latin '' bōs'': cow, ox, bull) is a genus of bovines, which includes, among others, wild and domestic cattle. ''Bos'' is often divided into four subgenera: ''Bos'', ''Bibos'', ''Novibos'', and ''Poephagus'', but including t ...
'' (cattle and yaks), ''
Ovis ''Ovis'' is a genus of mammals, part of the Caprinae subfamily of the ruminant family (biology), family Bovidae. Its seven highly sociable species are known as sheep or ovines. Domestic sheep are members of the genus, and are thought to be des ...
'' (sheep) and '' Capra'' (goats) about 36 million years ago. The Eurasian
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of Odocoileinae (Capreolini, Hydropotini and Alcini) split from the New World tribes of Capreolinae ( Odocoileini and Rangiferini) in the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
, 8.7–9.6 million years ago. ''Rangifer'' "evolved as a mountain deer, ...exploiting the subalpine and alpine meadows...". ''Rangifer'' originated in the
Late Pliocene Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
and diversified in the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
, a 2+ million-year period of multiple glacier advances and retreats. Several named ''Rangifer'' fossils in
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
and North America predate the evolution of modern tundra reindeer. Archaeologists distinguish "modern" tundra reindeer and barren-ground caribou from primitive forms – living and extinct – that did not have adaptations to extreme cold and to long-distance migration. They include a broad, high muzzle to increase the volume of the nasal cavity to warm and moisten the air before it enters the throat and lungs, bez tines set close to the brow tines, distinctive coat patterns, short legs and other adaptations for running long distances, and multiple behaviors suited to tundra, but not to forest (such as synchronized calving and aggregation during rutting and post-calving). As well, many genes, including those for
vitamin D Vitamin D is a group of structurally related, fat-soluble compounds responsible for increasing intestinal absorption of calcium, magnesium, and phosphate, along with numerous other biological functions. In humans, the most important compo ...
metabolism, fat metabolism, retinal development,
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural oscillation that repeats roughly every 24 hours. Circadian rhythms can refer to any process that originates within an organism (i.e., Endogeny (biology), endogenous) and responds to the env ...
, 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 In physical geography, a tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. There are three regions and associated types of tundra: #Arctic, Arctic, Alpine tundra, Alpine, and #Antarctic ...
or
polar desert Polar deserts are the regions of Earth that fall under an ice cap climate (''EF'' under the Köppen classification). Despite rainfall totals low enough to normally classify as a desert, polar deserts are distinguished from true deserts (' or ' un ...
s. The oldest undoubted ''Rangifer'' fossil is from
Omsk Omsk (; , ) is the administrative center and largest types of inhabited localities in Russia, city of Omsk Oblast, Russia. It is situated in southwestern Siberia and has a population of over one million. Omsk is the third List of cities and tow ...
, Russia, dated to 2.1-1.8 Ma. The oldest North American ''Rangifer'' fossil is from the
Yukon Yukon () is a Provinces and territories of Canada, territory of Canada, bordering British Columbia to the south, the Northwest Territories to the east, the Beaufort Sea to the north, and the U.S. state of Alaska to the west. It is Canada’s we ...
, 1.6 million years before present (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 The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
(Günz) Period, 680,000-620,000 BP. ''Rangifer'' fossils become increasingly frequent in circumpolar deposits beginning with the
Riss glaciation The Riss glaciation, Riss Glaciation, Riss ice age, Riss Ice Age, Riss glacial or Riss Glacial (, ', ' or (obsolete) ') is the second youngest glaciation of the Pleistocene epoch in the traditional, quadripartite glacial classification of the Alps ...
s, the second youngest of the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
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 Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...
that predate the
Last Glacial Maximum The Last Glacial Maximum (LGM), also referred to as the Last Glacial Coldest Period, was the most recent time during the Last Glacial Period where ice sheets were at their greatest extent between 26,000 and 20,000 years ago. Ice sheets covered m ...
(LGM) are of
Rancholabrean The Rancholabrean North American Land Mammal Age on the geologic timescale is a North American faunal stage in the North American Land Mammal Ages chronology (NALMA),Sanders, A.E., R.E. Weems, and L.B. Albright III (2009) Formalization of the mid- ...
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 Alabama ( ) is a U.S. state, state in the Southeastern United States, Southeastern and Deep South, Deep Southern regions of the United States. It borders Tennessee to the north, Georgia (U.S. state), Georgia to the east, Florida and the Gu ...
; and in
Sangamonian The Sangamonian Stage (or Sangamon interglacial) is the term used in North America to designate the Last Interglacial (130,000-115,000 years ago) and depending on definition, part of the early Last Glacial Period, corresponding to Marine Isotope St ...
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. The late Valerius Geist (1998) dates the Eurasian reindeer radiation dates to the large Riss glaciation (347,000 to 128,000 years ago), based on the Norwegian-Svalbard split 225,000 years ago. Finnish forest reindeer (''R. t. fennicus'') likely evolved from ''Cervus angifergeuttardi'' 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
Weichsel glaciation Weichsel may refer to: * Vistula river (Weichsel in German) * Weichselian glaciation * Peter Weichsel Peter M. Weichsel (born 1943) is an American professional bridge player from Encinitas, California. College and war years Early Weichsel st ...
);. The fossil species ''geuttardi'' was later replaced by ''R. constantini'', which was adapted for grasslands, 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. ''R. constantini'' was then replaced by modern tundra / barren-ground caribou adapted to extreme cold, probably in Beringia, before dispersing west (''R. t. tarandus'' in the
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
n mountains and ''R. t. sibiricus'' across Siberia) and east (''R. t. arcticus'' in the North American Barrenlands) when rising seas isolated them. Likewise in North America,
DNA analysis Genetic testing, also known as DNA testing, is used to identify changes in DNA sequence or chromosome structure. Genetic testing can also include measuring the results of genetic changes, such as RNA analysis as an output of gene expression, or ...
shows that woodland caribou (''R. caribou'') diverged from primitive ancestors of tundra / barren-ground caribou not during the LGM, 26,000–19,000 years ago, as previously assumed, but in the Middle Pleistocene around 357,000 years ago. 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 angiferhypogaeus'', 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 The Mesolithic (Ancient Greek language, Greek: μέσος, ''mesos'' 'middle' + λίθος, ''lithos'' 'stone') or Middle Stone Age is the Old World archaeological period between the Upper Paleolithic and the Neolithic. The term Epipaleolithic i ...
and
Neolithic The Neolithic or New Stone Age (from Ancient Greek, Greek 'new' and 'stone') is an archaeological period, the final division of the Stone Age in Mesopotamia, Asia, Europe and Africa (c. 10,000 BCE to c. 2,000 BCE). It saw the Neolithic Revo ...
Periods, and humans are today the main predator in many areas.
Norway Norway, officially the Kingdom of Norway, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. The remote Arctic island of Jan Mayen and the archipelago of Svalbard also form part of the Kingdom of ...
and
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
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 Jotunheimen (; "the home of the Jötunn") is a mountainous area of roughly in southern Norway and is part of the long range known as the Scandinavian Mountains. The 29 highest mountains in Norway are all located in the Jotunheimen mountains, in ...
, it is still possible to find remains of stone-built
trapping pit Trapping pits are deep pits dug into the ground, or built from stone, in order to trap animals. European rock drawings and cave paintings reveal that bear, moose and wolf were hunted since the Stone Age using trapping pits. Remains of trapping ...
s, guiding fences and bow rests, built especially for hunting reindeer. These can, with some certainty, be dated to the
Migration Period The Migration Period ( 300 to 600 AD), also known as the Barbarian Invasions, was a period in European history marked by large-scale migrations that saw the fall of the Western Roman Empire and subsequent settlement of its former territories ...
, although it is not unlikely that they have been in use since the
Stone Age The Stone Age was a broad prehistory, prehistoric period during which Rock (geology), stone was widely used to make stone tools with an edge, a point, or a percussion surface. The period lasted for roughly 3.4 million years and ended b ...
.
Cave painting In archaeology, cave paintings are a type of parietal art (which category also includes petroglyphs, or engravings), found on the wall or ceilings of caves. The term usually implies prehistoric art, prehistoric origin. These paintings were often c ...
s by ancient Europeans include both tundra and forest types of reindeer. A 2022 study of ancient
environmental DNA Environmental DNA or eDNA is DNA that is collected from a variety of environmental samples such as soil, seawater, snow or air, rather than directly sampled from an individual organism. As various organisms interact with the environment, DNA ...
from the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
(2 million years ago) Kap Kobenhavn Formation of northern Greenland identified preserved DNA fragments of ''Rangifer'', identified as basal but potentially ancestral to modern reindeer. This suggests that reindeer have inhabited Greenland since at least the Early Pleistocene. Around this time, northern Greenland was warmer than the
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
, with a
boreal forest Taiga or tayga ( ; , ), also known as boreal forest or snow forest, is a biome characterized by pinophyta, coniferous forests consisting mostly of pines, spruces, and larches. The taiga, or boreal forest, is the world's largest land biome. I ...
hosting a species assemblage with no modern analogue. These are among the oldest DNA fragments ever sequenced.


Taxonomy


Naming and research on museum collections

Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in 1758 named the Eurasian tundra species ''Cervus tarandus'', the genus ''Rangifer'' being credited to Smith, 1827. ''Rangifer'' has had a convoluted history because of the similarity in antler architecture (brow tines asymmetrical and often palmate, bez tines, a back tine sometimes branched, and branched at the distal end, often palmate). Because of individual variability, early taxonomists were unable to discern consistent patterns among populations, nor could they, examining collections in Europe, appreciate the difference in habitats and the differing function they imposed on antler architecture. Comparative morphometrics, the measurement of skulls, is often seen as more objective than description of differences of color or antler patterns, but actually confounds genetic variance with epistatic and statistical variance as well as compounded environment-based variance. For example, woodland caribou males, rutting in boreal forest where only a few females can be found, collect harems and defend them against other males, for which they have short, straight, strong, much-branched antlers, beams flattened in cross-section, designed for combat — and not too large, so as not to impede them in forested winter ranges. By contrast, modern tundra caribou (see
Evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
above) have synchronized calving as a predator-avoidance strategy, which requires large rutting aggregations. Males cannot defend a harem because, while he was busy fighting, they would disappear into the mass of the herd. Males therefore tend individual females; their fights are infrequent and brief. Their antlers are thin, beams round in cross-section, sweep back and then forward with a cluster of branches at the top; these are designed more for visual stimulation of the females. Their bez tines are set low, just above the brow tine, which is vertically flattened to protect the eyes while the buck "threshes" low brush, a courtship display. 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.


Biological exploration expeditions

Towards the end of the 19th century, national museums began sending out biological exploration expeditions and collections accumulated. Taxonomists, usually working for the museums, began naming subspecies more rigorously, based on statistical differences in detailed cranial, dental and skeletal measurements than antlers and pelage, supplemented by better knowledge of differences in ecology and behavior. From 1898 to 1937,
mammalogist In zoology, mammalogy is the study of mammals – a class of vertebrates with characteristics such as homeothermic metabolism, fur, four-chambered hearts, and complex nervous systems. The archive of number of mammals on earth is constantly growin ...
s named 12 new species (other than barren-ground and woodland, which had been named earlier) of caribou in Canada and Alaska, and three new species and nine new subspecies in Eurasia, each properly described according to the evolving rules of zoological nomenclature, with type localities designated and type specimens deposited in museums (see table in Species and subspecies below). Supplementary file 1 for .


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 Labrador () is a geographic and cultural region within the Canadian province of Newfoundland and Labrador. It is the primarily continental portion of the province and constitutes 71% of the province's area but is home to only 6% of its populatio ...
caribou), ''R. t. osborni'' (Osborn's caribou — from
British Columbia British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...
) and ''R. t. terranovae'' (the
Newfoundland Newfoundland and Labrador is the easternmost province of Canada, in the country's Atlantic region. The province comprises the island of Newfoundland and the continental region of Labrador, having a total size of . As of 2025 the population ...
caribou) as invalid and included only
barren-ground caribou The barren-ground caribou (''Rangifer tarandus groenlandicus''; but subject to a recent taxonomic revision) is a subspecies of the reindeer (or the caribou in North America) that is found in the Canadian territories of Nunavut and the Northwest ...
, 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 Ian McTaggart-Cowan (June 25, 1910 – April 18, 2010) was a Scottish-Canadian zoologist, conservationist, and television presenter. He has been called "the father of Canadian ecology". He was the brother of meteorologist Patrick McTaggart-Cow ...
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'', referenced by the
American Society of Mammalogists The American Society of Mammalogists (ASM) was founded in 1919. Its primary purpose is to encourage the study of mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence ...
, English zoologist Peter Grubb agreed with Valerius Geist, a specialist on large mammals, that these subspecies were valid (i.e., before the recent revision): In North America, ''R. t. caboti'', ''R. t. caribou'', ''R. t. dawsoni'', ''R. t. groenlandicus'', ''R. t. osborni'', ''R. t. pearyi'', and ''R. t. terranovae''; and in Eurasia, ''R. t. tarandus'', ''R. t. buskensis'' (called ''R. t. valentinae'' in Europe; see below), ''R. t. phylarchus'', ''R. t. pearsoni'', ''R. t. sibiricus'' and ''R. t. platyrhynchus''. These subspecies were retained in the 2011 replacement work ''
Handbook of the Mammals of the World ''Handbook of the Mammals of the World'' (''HMW'') is a book series from the publisher Lynx Edicions. The nine volumes were published from 2009 to 2019. Each mammal family is assessed in a full text introduction with photographs and each species ...
Vol. 2: Hoofed Mammals''. Most Russian authors also recognized ''R. t. angustirostris'', a forest reindeer from east of
Lake Baikal Lake Baikal is a rift lake and the deepest lake in the world. It is situated in southern Siberia, Russia between the Federal subjects of Russia, federal subjects of Irkutsk Oblast, Irkutsk Oblasts of Russia, Oblast to the northwest and the Repu ...
. However, since 1991, many genetic studies have revealed deep divergence between modern tundra reindeer and woodland caribou.Røed, K.H.; Feruson, M.D.; Crête, M.; Bergerud, A.T. (1991) Genetic variation in transferrin as a predictor for differentiation and evolution of caribou from
eastern Canada Eastern Canada (, also the Eastern provinces, Canadian East or the East) is generally considered to be the region of Canada south of Hudson Bay/ Hudson Strait and east of Manitoba, consisting of the following provinces (from east to west): Newf ...
. Rangifer 11: 65-74.
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 image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
on subspecies, designations based on
ecotype Ecotypes are organisms which belong to the same species but possess different phenotypical features as a result of environmental factors such as elevation, climate and predation. Ecotypes can be seen in wide geographical distributions and may event ...
s, 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, an adaptation of "evolutionary significant units". The 12 designatable units for caribou in Canada (that is, excluding Alaska and Greenland) based on ecology, behavior and, importantly, genetics (but excluding
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
and archaeology) essentially followed the previously named subspecies distributions, without naming them as such, plus some ecotypes. Ecotypes are not
phylogenetically In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
based and cannot substitute for taxonomy.


Genetic, molecular, and archaeological evidence

Meanwhile,
genetic data A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
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 The Svalbard reindeer (''Rangifer tarandus platyrhynchus'') is a small subspecies or species of reindeer found on the Svalbard archipelago of Norway. Males average in weight, females , while for other reindeer generally body mass is for males a ...
(''R. t. platyrhynchus''), although not closely related to each other, were the most genetically divergent among ''Rangifer'' clades; that modern (see
Evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
above) Eurasian tundra reindeer (''R. t. tarandus'' and ''R. t. sibiricus'') and North American barren-ground caribou (''R. t. arcticus''), although sharing ancestry, were separable at the subspecies level; that Finnish forest reindeer (''R. t. fennicus'') clustered well apart from both wild and domestic tundra reindeer and that boreal woodland caribou (''R. t. caribou'') were separable from all others. Meanwhile, archaeological evidence was accumulating that Eurasian forest reindeer descended from an extinct forest-adapted reindeer and not from tundra reindeer (see
Evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
above); since they do not share a direct
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. According to modern evolutionary biology, all living beings could be descendants of a unique ancestor commonl ...
, they cannot be
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...
. 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 Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
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 > 60,000 years agoYannic, G.; Pellissier, L.; Ortego, J.; Lecomte, N.; Couturier, S.; Cuyler, C.; Dussault, C.; Hundertmark, K.J.; Irvine, R.J.; Jenkins, D.A.; Kolpashikov, L.; Mager, K.; Musiani, M.; Parker, K.L.; Røed, K.H.; Sipko, T.; Þórisson, S.G.; V.Weckworth, B.; Guisan, A.; Bernatchez, L.; Côté, S.D. (2013) Genetic diversity in caribou linked to past and future climate change. Nature Climate Change 4: 132-137. — 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 1909Osgood, W.H. (1909) Biological investigations in Alaska and Yukon Territory. US Department of Agriculture Biological survey of North American fauna 1: 1-285. Murie, 1935 and Anderson 1946, among others) ''R. t. stonei'', ''R. t. montanus'', ''R. t. fortidens'' and ''R. t. osborni'', respectively, and this phylogeny was confirmed by genetic analysis.


Novel genetics-based clades

DNA also revealed three unnamed clades that, based on genetic distance, genetic divergence and shared vs. private
haplotype A haplotype (haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material (DNA) which is inherited from two parents. Normally these organisms have their DNA orga ...
s and
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
s, together with ecological and behavioral differences, may justify separation at the subspecies level: the Atlantic- Gaspésie caribou (COSEWIC DU11), an eastern montane ecotype of the boreal woodland caribou, and the
Baffin Island Baffin Island (formerly Baffin Land), in the Canadian territory of Nunavut, is the largest island in Canada, the second-largest island in the Americas (behind Greenland), and the fifth-largest island in the world. Its area is (slightly smal ...
caribou. 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 Reindeer distribution#Ahiak, Beverly and Qamanirjuaq caribou herds, 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

Abbreviations: *''AMNH'' - American Museum of Natural History *''BCPM'' - British Columbia Provincial Museum (= ''RBCM'' the Royal British Columbia Museum) *''NHMUK'' - British Museum (Natural History) (originally the ''BMNH'') *''DMNH'' - Denver Museum of Nature and Science, Denver Museum of Natural History *''MCZ'' - Museum of Comparative Zoology *''MSI'' - Museum of the Smithsonian Institution *''NMC'' - National Museum of Canada (originally the ''CGS'' Canadian Geological Survey Museum, now the ''CMN'' Canadian Museum of Nature) *''NR'' - Swedish Museum of Natural History, Naturhistoriska Riksmuseet *''RSMNH'' - Royal Swedish Museum of Natural History *''USNM'', - Smithsonian Institution, United States National Museum *''ZMASL'' - Zoological Museum (Saint Petersburg), 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; and four mountain ecotypes now known to be of distant
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...
-
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
lineage (see Reindeer#Taxonomy, Taxonomy above). The scientific name ''Tarandus rangifer buskensis'' Millais, 1915 (the Busk Mountains reindeer) was selected as the senior synonym to ''R. t. valentinae'' Flerov, 1933, in ''Mammal Species of the World'' but Russian authors do not recognize Millais and Millais' articles in a hunting travelogue, ''The Gun at Home and Abroad'',Millais, J.G. (1915) The Asiatic reindeer and elk. In: Carruthers, D.; Millais, J.G.; Wallace, H.F.; Kennion, L.C.R.L.; Barklay, F.G. (eds.) ''The Gun at Home and Abroad''. London & Counties Press Association Ltd., London, U.K., 216-223. seem short of a taxonomic authority. The scientific name ''groenlandicus'' is fraught with problems. Edwards (1743) 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) in his famously flawed revision, gave the name ''groenlandicus'' to all the barren-ground caribou in North America, Greenland included, because ''groenlandicus'' pre-dates Richardson's ''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%, whereas most cervid (deer family) species have a genetic distance of 2% to 5%)--as well as behavioral and morphological differences—a recent revision returned it to species status as ''R. groenlandicus''. Although it has been assumed that the larger caribou that appeared in Greenland 4,000 years ago originated from
Baffin Island Baffin Island (formerly Baffin Land), in the Canadian territory of Nunavut, is the largest island in Canada, the second-largest island in the Americas (behind Greenland), and the fifth-largest island in the world. Its area is (slightly smal ...
(itself unique; see Reindeer#Taxonomy, Taxonomy above), a reconstruction of LGM glacial retreat and caribou advance (Yannic et al. 2013) shows colonization by NAL lineage caribou more likely. Their PCA and tree diagrams show Greenland caribou clustering outside of the Beringian-Eurasian lineage. The scientific name ''R. t. granti'' has a very interesting history. Allen (1902) named it as a distinct species, ''R. granti'', from the "western end of
Alaska Peninsula The Alaska Peninsula (also called Aleut Peninsula or Aleutian Peninsula, ; Sugpiaq language, Sugpiaq: ''Aluuwiq'', ''Al'uwiq'') is a peninsula extending about to the southwest from the mainland of Alaska and ending in the Aleutian Islands. T ...
, opposite Popof Island, Popoff Island" and noting that:
''Rangifer granti'' is a representative of the Barren Ground group of Caribou, which includes ''R. arcticus'' of the Arctic Coast and ''R. granlandicus'' of Greenland. It is not closely related to ''R. stonei'' of the Kenai Peninsula, from which it differs not only in its very much smaller size, but in important cranial characters and in coloration. ...The external and cranial differences between ''R. granti'' and the various forms of the Woodland Caribou are so great in almost every respect that no detailed comparison is necessary. ...According to Mr. Stone, ''Rangifer granti'' inhabits the " barren land of Alaska Peninsula, ranging well up into the mountains in summer, but descending to the lower levels in winter, generally feeding on the low flat lands near the coast and in the foothills...As regards cranial characters no comparison is necessary with ''R. montanus'' or with any of the woodland forms."
Osgood and Murie (1935), agreeing with ''granti''s close relationship with the barren-ground caribou, brought it under ''R. arcticus'' as a subspecies, ''R. t. granti''. Anderson (1946) and Banfield (1961), based on statistical analysis of cranial, dental and other characters, agreed. But Banfield (1961) also synonymized Alaska's large ''R. stonei'' with other mountain caribou of British Columbia and the Yukon as invalid subspecies of woodland caribou, then ''R. t. caribou''. This left the small, migratory barren-ground caribou of Alaska and the Yukon, including the
Porcupine caribou The Porcupine caribou is a herd or ecotype of the mainland barren-ground caribou (''Rangifer arcticus arcticus'', Synonym (taxonomy), syn. ''R. tarandus groenlandicus''Harding LE (2022) Available names for Rangifer (Mammalia, Artiodactyla, Cervid ...
herd, without a name, which Banfield rectified in his 1974 ''Mammals of Canada'' by extending to them the name "''granti''". The late 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 The Alaska Peninsula (also called Aleut Peninsula or Aleutian Peninsula, ; Sugpiaq language, Sugpiaq: ''Aluuwiq'', ''Al'uwiq'') is a peninsula extending about to the southwest from the mainland of Alaska and ending in the Aleutian Islands. T ...
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''), a large montane type, was described from the Kenai Peninsula (where, apparently, it was never common except in years of great abundance), the eastern end of the Alaska Peninsula, and mountains throughout southern and eastern Alaska.Allen, J.A. (1901) Description of a new caribou from Kenai Peninsula, Alaska. Bulletin of the American Museum of Natural History XIV: 143-148. It was placed under ''R. arcticus'' as a subspecies, ''R. t. stonei'', and later synonymised as noted above. The same genetic analyses mentioned above for ''R. t. granti'' resulted in resurrecting ''R. t. stonei'' as well. The Sakhalin reindeer (''R. t. setoni''), endemic to
Sakhalin Sakhalin ( rus, Сахали́н, p=səxɐˈlʲin) is an island in Northeast Asia. Its north coast lies off the southeastern coast of Khabarovsk Krai in Russia, while its southern tip lies north of the Japanese island of Hokkaido. An islan ...
, was described as ''Rangifer tarandus setoni'' Flerov, 1933, but Banfield (1961) brought it under ''R. t. fennicus'' as a junior synonym. The wild reindeer on the island are apparently extinct, having been replaced by domestic reindeer. Some of the ''Rangifer'' species and subspecies may be further divided by
ecotype Ecotypes are organisms which belong to the same species but possess different phenotypical features as a result of environmental factors such as elevation, climate and predation. Ecotypes can be seen in wide geographical distributions and may event ...
depending on several behavioral factors – predominant habitat use (northern, tundra, mountain, forest, boreal forest, forest-dwelling, woodland, woodland (boreal), woodland (migratory) or woodland (mountain), spacing (dispersed or aggregated) and migration patterns (sedentary or migratory). North American examples of this are the Torngat Mountains, 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 Baffin Island (formerly Baffin Land), in the Canadian territory of Nunavut, is the largest island in Canada, the second-largest island in the Americas (behind Greenland), and the fifth-largest island in the world. Its area is (slightly smal ...
caribou, an ecotype of the barren-ground caribou (''R. t. arcticus''); and the Dolphin-Union caribou, Dolphin-Union "herd", another ecotype of ''R. t. arcticus''. The last three of these likely qualify as subspecies, but they have not yet been formally described or named.


Physical characteristics

Naming in this and following sections follows the taxonomy in the authoritative 2011 reference work ''
Handbook of the Mammals of the World ''Handbook of the Mammals of the World'' (''HMW'') is a book series from the publisher Lynx Edicions. The nine volumes were published from 2009 to 2019. Each mammal family is assessed in a full text introduction with photographs and each species ...
Vol. 2: Hoofed Mammals''.


Antlers

In most Deer, cervid species, only males grow
antler Antlers are extensions of an animal's skull found in members of the Cervidae (deer) Family (biology), family. Antlers are a single structure composed of bone, cartilage, fibrous tissue, skin, nerves, and blood vessels. They are generally fo ...
s; the reindeer is the only cervid species in which females also grow them normally. 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), but on average the bull's antlers are the second largest of any extant deer, after those of the male moose. In the largest subspecies, the antlers of large bulls can range up to in width and in beam length. They have the largest antlers relative to body size among living deer species. Antler size measured in number of points reflects the nutritional status of the reindeer and climate variation of its environment. The number of points on male reindeer increases from birth to 5 years of age and remains relatively constant from then on. "In male caribou, antler mass (but not the number of tines) varies in concert with body mass." 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. Geist describes them as frontally emphasized, flat-beamed antlers. Woodland caribou antlers are thicker and broader than those of the barren-ground caribou and their legs and heads are longer. Quebec-Labrador male caribou antlers can be significantly larger and wider than other woodland caribou. Central barren-ground male caribou antlers are perhaps the most diverse in configuration and can grow to be very high and wide. Osborn's caribou antlers are typically the most massive, with the largest circumference measurements.Allen, J.A. (1902) Description of a new caribou from northern British Columbia and remarks on ''Rangifer montanus''. Bulletin of the American Museum of Natural History XVI: 149-158. 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 antler, velvet, filled with blood vessels and spongy in texture. The antler velvet of the
barren-ground caribou The barren-ground caribou (''Rangifer tarandus groenlandicus''; but subject to a recent taxonomic revision) is a subspecies of the reindeer (or the caribou in North America) that is found in the Canadian territories of Nunavut and the Northwest ...
and the
boreal woodland caribou The boreal woodland caribou (''Rangifer tarandus caribou''; but subject to a recent taxonomic revision. See Reindeer: Taxonomy), also known as Eastern woodland caribou, boreal forest caribou and forest-dwelling caribou, is a North American subsp ...
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 Inuit (singular: Inuk) are a group of culturally and historically similar Indigenous peoples traditionally inhabiting the Arctic and Subarctic regions of North America and Russia, including Greenland, Labrador, Quebec, Nunavut, the Northwe ...
, 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." Male reindeer use their antlers to compete with other males during the mating season. Butler (1986) showed that the social requirements of caribou females during the rut determines the mating strategies of males and, consequently, the form of male 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
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
n 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. Cows in good nutritional condition, for example, during a mild winter with good winter range quality, may grow new antlers earlier as antler growth requires high intake. According to a respected Igloolik elder, Noah Piugaattuk, who was one of the last outpost camp leaders, caribou (''tuktu'') antlers According to the Igloolik Oral History Project (IOHP), "Caribou antlers provided the Inuit with a myriad of implements, from Snow knife, 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". Currently, the larger racks of antlers are used by Inuit in Inuit art as materials for carving. Iqaluit-based Jackoposie Oopakak's 1989 carving, entitled ''Nunali'', which means "place where people live", and which is part of the permanent collection of the National Gallery of Canada, includes a massive set of caribou antlers on which he has intricately carved the miniaturized world of the Inuit where "Arctic birds, caribou, polar bears, seals, and whales are interspersed with human activities of fishing, hunting, cleaning skins, stretching boots, and travelling by dog sled and kayak...from the base of the antlers to the tip of each branch".


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 The Peary caribou (''Rangifer arcticus pearyi'') is a subspecies of caribou found in the Canadian high Arctic islands of Nunavut and the Northwest Territories in Canada. They are the smallest of the North American caribou, with the females wei ...
, is the whitest and smallest subspecies of the continent, while the Selkirk Mountains caribou (Southern Mountain population DU9) is the darkest and nearly the largest,Reid, F. (2006). ''Mammals of North America''. Peterson Field Guides. only exceeded in size by Osborn's caribou (Northern Mountain population DU7). The Animal coat, coat has two layers of fur: a dense woolly undercoat and a longer-haired overcoat consisting of hollow, air-filled hairs.{{efn, According to Inuit elder Marie Kilunik of the Aivilingmiut, Canadian Inuit preferred the caribou skins from caribou taken in the late summer or autumn, when their coats had thickened. They used it for winter clothing "because each hair is hollow and fills with air trapping heat." Fur is the primary insulation factor that allows reindeer to regulate their Human body temperature#Core temperature, core body temperature in relation to their environment, the Temperature gradient, thermogradient, even if the temperature rises to {{cvt, 100, F, order=flip.{{cite journal , last=Moote , first=I. , year=1955 , title=The thermal insulation of caribou pelts , journal=Textile Research Journal , volume=25 , number =10 , pages=832–837 , doi=10.1177/004051755502501002 , s2cid=138926309 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".{{citation , first=Arthur Radclyffe , last=Dugmore , title=The romance of the Newfoundland caribou, page=191 , year=1913 , access-date=2 November 2014 , location=Philadelphia , publisher=Lippincott , url=https://archive.org/stream/romanceofnewfoun00dugm/romanceofnewfoun00dugm_djvu.txt A darker belly color may be caused by two mutations of Melanocortin 1 receptor, MC1R. They appear to be more common in domestic reindeer herds.{{cite journal , title=Two Missense Mutations in Melanocortin 1 Receptor (MC1R) Are Strongly Associated With Dark Ventral Coat Color in Reindeer (''Rangifer Tarandus'') , year=2014 , doi=10.1111/age.12187 , hdl=2164/4960, hdl-access=free, last1=Våge , first1=D. I. , last2=Nieminen , first2=M. , last3=Anderson , first3=D. G. , last4=Røed , first4=K. H. , journal=Animal Genetics , volume=45 , issue=5 , pages=750–753 , pmid=25039753


Heat exchange

Blood moving into the legs is cooled by blood returning to the body in a Countercurrent exchange, 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 {{cvt, 30, C-change 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.{{cite web , first=Mohd Hezri Fazalul , last=Rahiman , url=http://hezrif.uitm.edu.my/ese652/51%20Heat%20Exchanger%20090907d.pdf , title=Heat exchanger , year=2009 , access-date=3 November 2014 , location=Malaysia , archive-url=https://web.archive.org/web/20131205011425/http://hezrif.uitm.edu.my/ese652/51%20Heat%20Exchanger%20090907d.pdf , archive-date=5 December 2013 Reindeer have specialized counter-current vascular heat exchange in their Nasal cavity, nasal passages. Temperature gradient along the nasal Mucous membrane, mucosa is under physiological control. Incoming cold air is warmed by Thermoregulation, 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.{{cite journal , last1=Blix , first1=A.S. , last2=Johnsen , first2=Helge Kreiitzer, pmid=6887057, pmc=1199219 , title=Aspects of nasal heat exchange in resting reindeer , journal=Journal of Physiology , volume=340 , year=1983 , pages=445–454 , doi=10.1113/jphysiol.1983.sp014772 Like moose, caribou have specialized noses featuring nasal Nasal concha, 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 (Species at Risk Act, SARA), woodland {{blockquote, "Caribou have large feet with four toes. In addition to two small ones, called "dew claws," they have two large, crescent-shaped toes that support most of their weight and serve as shovels when digging for food under snow. These large concave hooves offer stable support on wet, soggy ground and on crusty snow. The pads of the hoof change from a thick, fleshy shape in the summer to become hard and thin in the winter months, reducing the animal's exposure to the cold ground. Additional winter protection comes from the long hair between the "toes"; it covers the pads so the caribou walks only on the horny rim of the hooves.", source=SARA 2014 Reindeer Hoof, hooves adapt to the season: in the summer, when the tundra is soft and wet, the footpads become sponge-like and provide extra traction. In the winter, the pads shrink and tighten, exposing the rim of the hoof, which cuts into the ice and crusted snow to keep it from slipping. This also enables them to dig down (an activity known as "cratering") through the snow to their favourite food, a lichen known as reindeer lichen (''Cladonia rangiferina'').


Size

The females (or "cows" as they are often called) usually measure {{cvt, 162, -, 205, cm in length and weigh {{cvt, 80, -, 120, kg.Caribou at the Alaska Department of Fish & Game
. Adfg.state.ak.us. Retrieved on 16 September 2011.
The males (or "bulls" as they are often called) are typically larger (to an extent which varies between the different species and subspecies), measuring {{cvt, 180, -, 214, cm in length and usually weighing {{cvt, 159, -, 182, kg. Exceptionally large bulls have weighed as much as {{cvt, 318, kg. Weight varies drastically between the seasons, with bulls losing as much as 40% of their pre-rut weight.{{citation , title=The Natural History of Canadian Mammals , last=Naughton , first=Donna , pages=543, 562, 567 , publisher=Canadian Museum of Nature and University of Toronto Press , isbn=978-1-4426-4483-0 , year=2011 The shoulder height is usually {{cvt, 85, to, 150, cm, and the tail is {{cvt, 14, to, 20, cm long. The Svalbard reindeer, reindeer from Svalbard are the smallest of all. They are also relatively short-legged and may have a shoulder height of as little as {{cvt, 80, cm,{{cite web , last1=Aanes , first1=Ronny , date=2007 , title=Svalbard reindeer , url=http://npweb.npolar.no/english/arter/svalbardrein , website=Norwegian Polar Institute , archive-url=https://web.archive.org/web/20101222190226/http://npweb.npolar.no/english/arter/svalbardrein , archive-date=22 December 2010 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.Banfield, Alexander William Francis (1966) "The caribou", pp. 25–28 in ''The Unbelievable Land''. Smith, I.N. (ed.) Ottawa: Queen's Press, cited in {{Cite journal , pmid = 18986518 , year = 2008 , last1 = Bro-Jørgensen , first1 = J. , title = Knee-clicks and visual traits indicate fighting ability in eland antelopes: Multiple messages and back-up signals , journal = BMC Biology , volume = 6 , page = 47 , last2 = Dabelsteen , first2 = T. , doi = 10.1186/1741-7007-6-47 , pmc = 2596769 , doi-access = free 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 clicking sound made by reindeer as they walk is caused by small tendons slipping over bone protuberances (sesamoid bones) in their feet.{{citation, title=Hoofed Mammals of British Columbia , first=David , last=Shackleton , date= May 2013 , publisher=Royal BC Museum , isbn=978-0-7726-6638-3 , orig-year=1999{{citation, last=Banfield , first=Alexander William Francis, chapter=The caribou, title=The Unbelievable Land, editor-last=Smith, editor-first=I.N., location=Ottawa, publisher=Queen's Press, year=1966, pages=25–28 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.


Eyes

A study by researchers from University College London in 2011 revealed that reindeer can see light with wavelengths as short as 320 nm (i.e. in the ultraviolet range), considerably below the human threshold of 400 nm. It is thought that this ability helps them to survive in the Arctic, because many objects that blend into the landscape in light visible to humans, such as urine and fur, produce sharp contrasts in ultraviolet.Reindeer use UV light to survive in the wild
{{webarchive , url=https://web.archive.org/web/20111129042727/http://www.ucl.ac.uk/news/news-articles/1105/11052502 , date=29 November 2011 . Ucl.ac.uk (26 May 2011). Retrieved on 16 September 2011.
It has been proposed that Corona discharge#Problems, 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."{{cite journal , author1=Tyler, N. , author2=Stokkan, K.A. , author3=Hogg, C. , author4=Nellemann, C. , author5=Vistnes, A.I. , author6=Jeffery, G. , year=2014 , title=Ultraviolet vision and avoidance of power lines in birds and mammals , journal=Conservation Biology , volume=28 , issue=3 , pages=630–631 , doi=10.1111/cobi.12262, pmid=24621320 , pmc=4232876 , bibcode=2014ConBi..28..630T 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


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 {{cvt, 10, kg 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 {{cvt, 3, kg. After this, however, non-breeding females on average have a higher body fat mass than do breeding females.{{cite journal , last1 = Allaye Chan-McLeod , first1 = A.C. , last2 = White , first2 = R.G. , last3 = Russell , first3 = D.E. , year = 1999 , title = Comparative body composition strategies of breeding and nonbreeding female caribou , journal = Canadian Journal of Zoology , volume = 77 , issue = 12, pages = 1901–1907 , doi=10.1139/z99-169, bibcode = 1999CaJZ...77.1901C 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. Lichens are a crucial part of the reindeer diet; however, they are less prevalent in the diet of pregnant reindeer compared to non-pregnant individuals. The amount of lichen in a diet is found more in non-pregnant adult diets than pregnant individuals due to the lack of nutritional value. Although lichens are high in carbohydrates, they are lacking in essential proteins that vascular plants provide. The amount of lichen in a diet decreases in latitude, which results in nutritional stress being higher in areas with low lichen abundance.{{cite journal , last1=Joly , first1=K. , last2=Wasser , first2=S. K. , last3=Booth , first3=R. , date=10 June 2015 , title=Non-Invasive Assessment of the Interrelationships of Diet, Pregnancy Rate, Group Composition, and Physiological and Nutritional Stress of Barren-Ground Caribou in Late Winter , journal=PLOS ONE , volume=10 , issue=6 , at=e0127586 , issn=1932-6203 , doi=10.1371/journal.pone.0127586 , pmid=26061003 , pmc=4464525 , bibcode=2015PLoSO..1027586J , doi-access=free{{rp, 6 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 University of Tromsø, 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

{{Further, Rut (mammalian reproduction)#Cervidae Reindeer mate in late September to early November, and the gestation period is about 228–234 days.{{citation , title=Caribou , first=A.T. , last=Bergerud , url=http://www.thecanadianencyclopedia.ca/en/article/caribou/ , publisher=The Canadian Encyclopedia , date=29 April 2014 , access-date=3 September 2014 , url-status=live , archive-url=https://web.archive.org/web/20141207050757/http://www.thecanadianencyclopedia.ca/en/article/caribou/ , archive-date=7 December 2014 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.{{cite web, url=http://www.animalcorner.co.uk/wildlife/caribou.html , title=Caribou at Animal Corner , url-status=dead , archive-url=https://web.archive.org/web/20121029063613/http://www.animalcorner.co.uk/wildlife/caribou.html , archive-date=29 October 2012 To calve, "females travel to isolated, relatively predator-free areas such as islands in lakes, peatlands, lake-shores, or tundra." As females select the habitat for the birth of their calves, they are warier than males. Dugmore noted that, in their seasonal migrations, the herd follows a female for that reason. Newborns weigh on average {{cvt, 6, kg. In May or June, the calves are born. After 45 days, the calves are able to graze and forage, but continue suckling until the following autumn when they become independent from their mothers. Bulls live four years less than the cows, whose maximum longevity is about 17 years. Cows with a normal body size and who have had sufficient summer nutrition can begin breeding anytime between the ages of 1 and 3 years. When a cow has undergone nutritional stress, it is possible for her to not reproduce for the year. Dominant bulls, those with larger body size and antler racks, inseminate more than one cow a season.


Social structure, migration and range

Some populations of North American caribou; for example, many herds in the barren-ground caribou subspecies and some woodland caribou in District of Ungava, Ungava and northern
Labrador Labrador () is a geographic and cultural region within the Canadian province of Newfoundland and Labrador. It is the primarily continental portion of the province and constitutes 71% of the province's area but is home to only 6% of its populatio ...
, Animal migration, migrate the farthest of any terrestrial mammal, traveling up to {{cvt, 5000, km, -3 a year, and covering {{cvt, 1000000, km2, -5. Other North American populations, the boreal woodland caribou for example, are largely sedentary.{{Cite journal , pmid = 21227095 , year = 1988 , last1 = Bergerud , first1 = A. T. , title = Caribou, wolves and man , journal = Trends in Ecology & Evolution , volume = 3 , issue = 3 , pages = 68–72 , doi = 10.1016/0169-5347(88)90019-5 , bibcode = 1988TEcoE...3...68B The European populations are known to have shorter migrations. Island populations, such as the Novaya Zemlya and Svalbard reindeer and the Peary caribou, make local movements both within and among islands. Migrating reindeer can be negatively affected by Parasitism, parasite 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.{{cite journal, last1=Bartel, first1=Rebecca, last2=Oberhauser, first2=Karen , author-link2=Karen Oberhauser , last3=De Roode, first3=Jacob, last4=Atizer, first4=Sonya , title=Monarch butterfly migration and parasite transmission in eastern North America, journal=Ecology, date=February 2011, volume=92, issue=2 , pages=342–351, doi=10.1890/10-0489.1, pmid=21618914, pmc=7163749, bibcode=2011Ecol...92..342B , s2cid=9018584 Normally travelling about {{cvt, 19, -, 55, km a day while migrating, the caribou can run at speeds of {{cvt, 60, -, 80, km/h. Young calves can already outrun an Olympic sprinter when only 1 day old.{{cite book, last=Hoare, first=Ben, title=Animal Migration, year=2009, publisher=Natural History Museum, location=London , isbn=978-0-565-09243-6, page=45 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 {{cvt, 6.5, km/h but, if necessary, at {{cvt, 10, km/h 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 caribou, Porcupine herd of barren-ground caribou are among the longest of any mammal. Greenland caribou, found in southwestern
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
, are "mixed migrators" and many individuals do not migrate; those that do migrate less than 60 km. 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. Although most wild tundra reindeer migrate between their winter range in taiga and summer range in tundra, some ecotypes or herds are more or less sedentary. Novaya Zemlya reindeer (''R. t. pearsoni'') formerly wintered on the mainland and migrated across the ice to the islands for summer, but only a few now migrate.{{Cite journal , last1=Mizin , first1=Ivan A. , last2=Sipko , first2=Taras P. , last3=Davydov , first3=Andrey V. , last4=Gruzdev , first4=Alexander R. , date=2018 , title=The wild reindeer (''Rangifer tarandus'': Cervidae, Mammalia) on the arctic islands of Russia: a review , journal=Nature Conservation Research , volume=3 , issue=3 , doi=10.24189/ncr.2018.040 , issn=2500-008X, doi-access=free 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. {{clear left As an adaptation to their Arctic environment, they have lost their
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural oscillation that repeats roughly every 24 hours. Circadian rhythms can refer to any process that originates within an organism (i.e., Endogeny (biology), endogenous) and responds to the env ...
. {{clear left


Ecology


Distribution and habitat

{{main, Reindeer distribution Originally, the reindeer was found in
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
, Eastern Europe,
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
, Russia, Mongolia and northern China north of the 50th parallel north, 50th latitude. In North America, it was found in Canada, Alaska, and the northern
contiguous United States The contiguous United States, also known as the U.S. mainland, officially referred to as the conterminous United States, consists of the 48 adjoining U.S. states and the District of Columbia of the United States in central North America. The te ...
from
Maine Maine ( ) is a U.S. state, state in the New England region of the United States, and the northeasternmost state in the Contiguous United States. It borders New Hampshire to the west, the Gulf of Maine to the southeast, and the Provinces and ...
to
Washington Washington most commonly refers to: * George Washington (1732–1799), the first president of the United States * Washington (state), a state in the Pacific Northwest of the United States * Washington, D.C., the capital of the United States ** A ...
. 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.{{cite news , last = Watson , first = Jeremy , date = 12 October 2006 , title = Sea eagle spreads its wings ... , location = Edinburgh , newspaper = Scotland on Sunday During the Late Pleistocene Epoch, reindeer occurred further south in North America, such as in Nevada, Tennessee, and
Alabama Alabama ( ) is a U.S. state, state in the Southeastern United States, Southeastern and Deep South, Deep Southern regions of the United States. It borders Tennessee to the north, Georgia (U.S. state), Georgia to the east, Florida and the Gu ...
, 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 Norway, officially the Kingdom of Norway, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. The remote Arctic island of Jan Mayen and the archipelago of Svalbard also form part of the Kingdom of ...
, 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, Sakhalin Island, and USA (northern Idaho and Great Lakes region)." Reindeer were introduced to, and are feral in, "Iceland, Kerguelen Islands, South Georgia, South Georgia Island, Pribilof Islands, St. Matthew Island"; a free-ranging semi-domesticated herd is also present in Scotland. 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 realm, Holarctic, being present in both
tundra In physical geography, a tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. There are three regions and associated types of tundra: #Arctic, Arctic, Alpine tundra, Alpine, and #Antarctic ...
and
taiga Taiga or tayga ( ; , ), also known as boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruces, and larches. The taiga, or boreal forest, is the world's largest land biome. In North A ...
(boreal forest),{{cite book , title=Walker's Mammals of the World , year=1999 , publisher=Johns Hopkins University Press , location=Baltimore , isbn=978-0-8018-5789-8 , pages=1128–1130 , edition=6th , volume=2 , editor=Novak, R. M. by 2013, many herds had "unusually low numbers" and their winter ranges in particular were smaller than they used to be. Caribou and reindeer numbers have fluctuated historically, but many herds are in decline across their range.{{cite web, url=http://news.bbc.co.uk/earth/hi/earth_news/newsid_8094000/8094036.stm , last=Walker, first=Matt , publisher=BBC , date=11 June 2009 , website=Earth News, title=Reindeer herds in global decline, archive-url=https://web.archive.org/web/20120103172521/http://news.bbc.co.uk/earth/hi/earth_news/newsid_8094000/8094036.stm , archive-date=3 January 2012 , access-date=16 September 2011, url-status=live This global decline is linked to climate change for northern migratory herds and industrial disturbance of habitat for non-migratory herds. {{cite journal , last1=Vors , first1=L. S. , last2=Boyce , first2=M. S. , year=2009 , title=Global declines of caribou and reindeer , journal=Global Change Biology , volume=15 , issue=11 , pages=2626–2633 , doi=10.1111/j.1365-2486.2009.01974.x , bibcode=2009GCBio..15.2626V , s2cid=86111815 , issn=1354-1013 Barren-ground caribou are susceptible to the effects of climate change due to a Match/mismatch, mismatch in the Phenology, phenological process between the availability of food during the calving period. In November 2016, it was reported that more than 81,000 reindeer in Russia had died as a result of climate change. Longer autumns, leading to increased amounts of freezing rain, created a few inches of ice over lichen, causing many reindeer to starve to death.


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 owing to specialised bacteria and protozoa in their gut.{{citation, title=Wolves in Canada, last=McCloskey, first=Erin, publisher=Lone Pines , year=2011 , chapter=Caribou , pages=72–82 , isbn=978-1-55105-872-6 They are also the only animals (except for some Gastropoda, gastropods) in which the enzyme lichenase, which breaks down lichenin to glucose, has been found. However, they also eat the leaves of willows and birches, as well as Cyperaceae, sedges and Poaceae, grasses. Reindeer are Osteophagy, 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 Egg, bird eggs. Reindeer herded by the Chukchi people, Chukchis have been known to devour mushrooms enthusiastically in late summer. During the Arctic summer, when there is Midnight sun, continuous daylight, reindeer change their sleeping pattern from one Circadian rhythm, synchronised with the sun to an Ultradian rhythm, ultradian pattern, in which they sleep when they need to digest food.{{cite web , url= https://www.livescience.com/62870-summer-solstice-animals.html , title=How Does the Summer Solstice Affect Animals? , last= Hickok , first= K. , date= 21 June 2018 , website= Live Science , access-date= 22 June 2018 Δ13C, ''δ''13CC values indicate reindeer living in the region around Biśnik Cave exhibited minimal ecological change during the transition from Marine isotope stages, 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. {{clear left


Predators

A variety of predators prey heavily on reindeer, including overhunting by people in some areas, which contributes to the decline of populations. Golden eagles prey on calves and are the most prolific hunter on the calving grounds. 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.{{cite journal , last1=McLoughlin , first1=P.D. , last2=Dzus , first2=E. , last3=Wynes , first3=B. , last4=Boutin , first4=Stan , year=2003 , title=Declines in populations of woodland caribou , journal=Journal of Wildlife Management , volume=67 , issue=4 , pages=755–761, doi= 10.2307/3802682 , jstor=3802682 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.{{cite web , last=Chapman , first=Andrew , date=October 22, 2021 , title=Polar bear hunting a reindeer caught on tape for first time. Unusual behavior had been assumed, but never clearly seen , website=Science (journal), Science , url=https://www.science.org/content/article/polar-bear-hunting-reindeer-caught-tape-first-time , access-date=October 26, 2021 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".{{cite journal , last1=Stempniewicz , first1=Lech , last2=Kulaszewicz , first2=Izabela , last3=Aars , first3=Jon , title=Yes, they can: polar bears ''Ursus maritimus'' successfully hunt Svalbard reindeer ''Rangifer tarandus platyrhynchus'' , journal=Polar Biology , publisher=Springer Science and Business Media LLC , volume=44 , issue=11 , date=October 12, 2021 , issn=0722-4060 , doi=10.1007/s00300-021-02954-w , pages=2199–2206, bibcode=2021PoBio..44.2199S , s2cid=241470816 , doi-access=free Additionally, as carrion, reindeer may be scavenged opportunistically by Red fox, red and Arctic foxes, various species of eagles, hawks and falcons, and common ravens. Bloodsucking insects, such as mosquitoes, Black fly, black flies, and especially the reindeer warble fly or reindeer botfly (''Hypoderma tarandi'') and the reindeer nose botfly (''Cephenemyia trompe''),{{cite journal , last=Cooper , first=Elisabeth J. , title=Warmer Shorter Winters Disrupt Arctic Terrestrial Ecosystems , journal=Annual Review of Ecology, Evolution, and Systematics , publisher=Annual Reviews (publisher), Annual Reviews , volume=45 , issue=1 , date=2014-11-23 , issn=1543-592X , doi=10.1146/annurev-ecolsys-120213-091620 , pages=271–295, doi-access=free 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 {{cvt, 1, L of blood to biting insects for every week it spends in the tundra. The population numbers of some of these predators is influenced by the migration of reindeer.{{Citation needed, date=May 2012 Tormenting insects keep caribou on the move, searching for windy areas like hilltops and mountain ridges, rock reefs, lakeshore and forest openings, or snow patches that offer respite from the buzzing horde. Gathering in large herds is another strategy that caribou use to block insects. 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{{cite journal , last=Duffy , first=Michael S. , author2=Nathan J. Keppie , author3=Michael D. B. Burt , title=Meningeal Worm is a Long-lived Parasitic Nematode in White-tailed Deer , journal=Journal of Wildlife Diseases , year=2002 , volume=38 , issue=2 , pages=448–452 , doi=10.7589/0090-3558-38.2.448 , pmid=12038147 , s2cid=39879199{{cite web , url=http://www.vet.utk.edu/news/story/brain-worm-%28meningeal-worm%29-infestation-in-llamas-and-alpacas.html , title="Brain Worm" (Meningeal Worm) Infestation in Llamas and Alpacas , publisher=University of Tennessee , access-date=14 November 2013, url-status=live , archive-url=https://web.archive.org/web/20131021025732/http://www.vet.utk.edu/news/story/brain-worm-%28meningeal-worm%29-infestation-in-llamas-and-alpacas.html , archive-date=21 October 2013 which include a loss of fear of humans. White-tailed deer that carry this worm are partially immune to it. Changes in climate and habitat beginning in the 20th century have expanded range overlap between white-tailed deer and caribou, increasing the frequency of infection within the reindeer population. This increase in infection is a concern for wildlife managers. Human activities, such as "clear-cutting forestry practices, forest fires, and the clearing for agriculture, roadways, railways, and power lines," favor the conversion of habitats into the preferred habitat of the white-tailed deer – "open forest interspersed with meadows, clearings, grasslands, and riparian flatlands." Towards the end of the Soviet Union, there was increasingly open admission from the Soviet government that reindeer numbers were being negatively affected by human activity, and that this must be remediated especially by supporting reindeer breeding by native herders.{{cite journal , last1=Astakhov , first1=Alexander S. , last2=Khaitun , first2=A. D. , last3=Subbotin , first3=G. E. , title=Socioeconomic Aspects of Oil and Gas Development in West Siberia , journal=Annual Review of Energy and the Environment , publisher=Annual Reviews (publisher), Annual Reviews , volume=14 , issue=1 , year=1989 , issn=0362-1626 , doi=10.1146/annurev.eg.14.110189.001001 , pages=117–130 , s2cid=154741203, doi-access=free


Conservation


Current status

According to the International Union for Conservation of Nature, 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 species, Vulnerable due to an observed population decline of 40% over the last +25 years. Some reindeer species and subspecies are rare and three subspecies have already become Extinction, extinct. In North America, the Queen Charlotte Islands caribou{{citation, url=http://www.cosewic.gc.ca/eng/sct4/result_e.cfm?SSGbox=All, year=2004, work=COSEWIC, institution=Committee on the Status of Endangered Wildlife in Canada, access-date=16 January 2014, title=Subcommittees, url-status=dead, archive-url=https://web.archive.org/web/20140201231054/http://www.cosewic.gc.ca/eng/sct4/result_e.cfm?SSGbox=All, archive-date=1 February 2014{{cite journal, author1=S. A. Byun , author2=B. F. Koop , author3=T. E. Reimchen , title=Evolution of the Dawson caribou (''Rangifer tarandus dawsoni''), journal= Can. J. Zool., volume= 80, issue=5, pages= 956–960, year=2002, doi=10.1139/z02-062, bibcode=2002CaJZ...80..956B , s2cid=4950388 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.{{citation, title=Migratory Tundra ''Rangifer'', first1=Don E., last1=Russell, first2=A., last2=Gunn, institution=NOAA Arctic Research Program, date=20 November 2013, access-date=14 January 2014, url=http://www.arctic.noaa.gov/reportcard/caribou_reindeer.html, series=Annual Arctic Report Card, url-status=live, archive-url=https://web.archive.org/web/20140121105443/http://www.arctic.noaa.gov/reportcard/caribou_reindeer.html, archive-date=21 January 2014 Grant's caribou, a small, pale subspecies endemic to the western end of the
Alaska Peninsula The Alaska Peninsula (also called Aleut Peninsula or Aleutian Peninsula, ; Sugpiaq language, Sugpiaq: ''Aluuwiq'', ''Al'uwiq'') is a peninsula extending about to the southwest from the mainland of Alaska and ending in the Aleutian Islands. T ...
and the adjacent islands, has not been assessed as to its conservation status. The status of the Dolphin-Union caribou, Dolphin-Union "herd" was upgraded to Endangered in 2017. 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 River, Columbia and Kootenay River, 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 Local extinction, extirpated in the
contiguous United States The contiguous United States, also known as the U.S. mainland, officially referred to as the conterminous United States, consists of the 48 adjoining U.S. states and the District of Columbia of the United States in central North America. The te ...
, including
Washington Washington most commonly refers to: * George Washington (1732–1799), the first president of the United States * Washington (state), a state in the Pacific Northwest of the United States * Washington, D.C., the capital of the United States ** A ...
and Idaho. Osborn's caribou (Northern Mountain population DU7) is classified as Threatened in Canada. In Eurasia, the Sakhalin reindeer is Extinction, 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. Many Siberian tundra reindeer herds have declined, some dangerously, but the Taymir herd remains strong and in total about 940,000 wild Siberian tundra reindeer were estimated in 2010. There is strong regional variation in ''Rangifer'' herd size. By 2013, many caribou herds in North America had "unusually low numbers" and their winter ranges in particular were smaller than they used to be. Caribou numbers have fluctuated historically, but many herds are in decline across their range. There are many factors contributing to the decline in numbers.


Boreal woodland caribou

Ongoing human development of their habitat has caused populations of boreal woodland caribou to disappear from their original southern range. In particular, boreal woodland caribou were Local extinction, extirpated in many areas of eastern North America in the beginning of the 20th century. Professor Marco Musiani of the University of Calgary said in a statement that "The woodland caribou is already an endangered subspecies in southern Canada and the United States...[The] warming of the planet means the disappearance of their critical habitat in these regions. Caribou need undisturbed lichen-rich environments and these types of habitats are disappearing." Boreal woodland caribou were designated as Threatened species, Threatened in 2002 by the
Committee on the Status of Endangered Wildlife in Canada The Committee on the Status of Endangered Wildlife in Canada (COSEWIC, French: Comité sur la situation des espèces en péril au Canada, COSEPAC) is an independent committee of wildlife experts and scientists whose "raison d'être is to identify s ...
, (COSEWIC). Environment Canada reported in 2011 that there were approximately 34 000 boreal woodland caribou in 51 ranges remaining in Canada (Environment and Climate Change 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 Peary caribou (''Rangifer arcticus pearyi'') is a subspecies of caribou found in the Canadian high Arctic islands of Nunavut and the Northwest Territories in Canada. They are the smallest of the North American caribou, with the females wei ...
. 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

{{See also, Reindeer in Siberian shamanism, Reindeer hunting in Greenland 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 The Finnish forest reindeer ''(Rangifer tarandus fennicus'' (Finnish:'' metsäpeura'', Russian: ''лесной северный олень''), also known as Eurasian or European forest reindeer is a rare subspecies of the reindeer native to Finla ...
or the narrow-nosed reindeer, an eastern Siberia forest form. Canadian examples include the Caribou Inuit, the inland-dwelling
Inuit Inuit (singular: Inuk) are a group of culturally and historically similar Indigenous peoples traditionally inhabiting the Arctic and Subarctic regions of North America and Russia, including Greenland, Labrador, Quebec, Nunavut, the Northwe ...
of the Kivalliq Region in northern Canada, the Caribou Clan in the Yukon, the Iñupiat, the
Inuvialuit The Inuvialuit (sing. Inuvialuk; ''the real people'') or Western Canadian Inuit are Inuit who live in the western Canadian Arctic region. They, like all other Inuit, are descendants of the Thule who migrated eastward from Alaska. Their homelan ...
, the Hän, the Northern Tutchone, and the
Gwichʼin The Gwichʼin (or Kutchin or Loucheux) are an Athabaskan languages, Athabaskan-speaking First Nations in Canada, First Nations people of Canada and an Alaskan Athabaskans, Alaska Native people. They live in the northwestern part of North America ...
(who followed the
Porcupine caribou The Porcupine caribou is a herd or ecotype of the mainland barren-ground caribou (''Rangifer arcticus arcticus'', Synonym (taxonomy), syn. ''R. tarandus groenlandicus''Harding LE (2022) Available names for Rangifer (Mammalia, Artiodactyla, Cervid ...
herd for millennia). Hunting wild reindeer and herding of semi-domesticated reindeer are important to several Arctic and sub-Arctic peoples such as the Dukha people, Duhalar for meat, {{Ill, Reindeer hide, lt=hides, de, Rentierfell, antlers, {{Ill, Reindeer milk, lt=milk, ru, Оленье молоко, and transportation."In North America and Eurasia the species has long been an important resource — in many areas ''the'' most important resource — for peoples inhabiting the northern boreal forest and tundra regions." (Banfield 1961:170; Kurtén 1968:170) {{cite journal , author=Ernest S. Burch Jr. , jstor=278435 , title=The Caribou/Wild Reindeer as a Human Resource , journal=American Antiquity , volume=37 , issue=3 , year=1972 , pages=339–368 , doi=10.2307/278435 , s2cid=161921691 Reindeer have been domesticated at least two and probably three times, in each case from wild Eurasian tundra reindeer after the Last Glacial Maximum, Last Glacial Maximum (LGM). Recognizably different domestic reindeer breeds include those of the Evenk, Even, and Chukotka-Khargin people of Yakutia and the Nenets breed from the Nenets Autonomous district and Murmansk region; 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 people, Sámi (Sápmi) have also depended on reindeer herding and fishing for centuries.{{citation , year=1971 , title=Atlas of Murmansk Oblast{{rp, IV{{citation , title=Administrative-Territorial Divisions of Murmansk Oblast{{rp, 16 In Sápmi, reindeer are used to pull a pulk, a Nordic sled. The reindeer has an important economic role for all circumpolar peoples, including the Sámi, the Swedes, the Norwegians, the Finns and the Northwestern Russians in Europe, the Nenets people, Nenets, the Khanty, the Evenks, the Yukaghir people, Yukaghirs, the Chukchi people, Chukchi and the Koryaks in Asia and the
Inuit Inuit (singular: Inuk) are a group of culturally and historically similar Indigenous peoples traditionally inhabiting the Arctic and Subarctic regions of North America and Russia, including Greenland, Labrador, Quebec, Nunavut, the Northwe ...
in North America. It is believed that domestication started between the Bronze Age, Bronze and Iron Ages. Siberian reindeer owners also use the reindeer to ride on (Siberian reindeer are larger than their Scandinavian relatives). For breeders, a single owner may own hundreds or even thousands of animals. The numbers of Russian and Scandinavian reindeer herders have been drastically reduced since 1990. The sale of fur and meat is an important source of income. Reindeer were introduced into Alaska near the end of the 19th century; they interbred with the native caribou subspecies there. Reindeer herders on the Seward Peninsula have experienced significant losses to their herds from animals (such as wolves) following the wild caribou during their migrations.{{Citation needed, date=May 2012 Reindeer meat is popular in the Scandinavian countries. Reindeer meatballs are sold canned. Sautéed reindeer is the best-known dish in Sápmi. In Alaska and Finland, reindeer sausage is sold in supermarkets and grocery stores. Reindeer meat is very tender and lean. It can be prepared fresh, but also dried, Salt-cured meat, salted and hot- and cold-Smoked meat, smoked. In addition to meat, almost all of the internal organs of reindeer can be eaten, some being traditional dishes. Furthermore, ''Lapin Poron liha'', fresh reindeer meat completely produced and packed in Finnish Lapland, is protected in Europe with Geographical indications and traditional specialities in the European Union#Protected designation of origin (PDO), PDO classification.Lapland Reindeer meat protected in the EU
{{webarchive , url=https://web.archive.org/web/20091207090514/http://60north.mmm.fi/news?&article=13673753§ion=05 , date=7 December 2009. North Magazine (Accessed 19 July 2010)

{{webarchive , url=https://web.archive.org/web/20100819142342/http://ec.europa.eu/agriculture/quality/door/list.html , date=19 August 2010. (Accessed 19 July 2010)
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 (drink), Caribou''.{{cite web, title=Quebec's Carnaval is worth freezing your a** off for , first=Julie , last=Ovenell-Carter , date=6 February 2009 , publisher=theseboots.travel , url=http://theseboots.travel/2009/02/06/worth-repeating-quebecs-carnaval-is-worth-freezing-your-a-off-for/ , url-status=usurped , archive-url=https://web.archive.org/web/20120303133943/http://theseboots.travel/2009/02/06/worth-repeating-quebecs-carnaval-is-worth-freezing-your-a-off-for/ , archive-date=3 March 2012


Indigenous North Americans

{{missing information, section, US government intervention to introduce herding in the form of Alaska Reindeer Service; Canadian purchase from Alaska, date=March 2023 Caribou are still hunted in Greenland and in North America. In the traditional lifestyles of some of Canada's
Inuit Inuit (singular: Inuk) are a group of culturally and historically similar Indigenous peoples traditionally inhabiting the Arctic and Subarctic regions of North America and Russia, including Greenland, Labrador, Quebec, Nunavut, the Northwe ...
peoples and northern First Nations in Canada, First Nations peoples, Alaska Natives, and the Kalaallit of Greenland, caribou is an important source of food, clothing, shelter and tools. The Caribou Inuit are inland-dwelling Inuit in present-day
Nunavut Nunavut is the largest and northernmost Provinces and territories of Canada#Territories, territory of Canada. It was separated officially from the Northwest Territories on April 1, 1999, via the ''Nunavut Act'' and the Nunavut Land Claims Agr ...
's Kivalliq Region (formerly the Keewatin Region, Northwest Territories), 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: {{blockquote, The caribou feeds the wolf, but it is the wolf who keeps the caribou strong., Kivalliq region Elder Chief of Koyukuk and chair for the Western Arctic Caribou Herd Working Group Benedict Jones, or Kʼughtoʼoodenoolʼoʼ, represents the Middle Yukon River, Alaska. His grandmother was a member of the Caribou Clan, who travelled with the caribou as a means to survive. In 1939, they were living their traditional lifestyle at one of their hunting camps in Koyukuk near the location of what is now the Koyukuk National Wildlife Refuge. His grandmother made a pair of new mukluks in one day. Kʼughtoʼoodenoolʼoʼ recounted a story told by an elder, who "worked on the steamboats during the gold rush days out on the Yukon." In late August, the caribou migrated from the Alaska Range up north to Huslia, Alaska, Huslia, Koyukuk and the Tanana, Alaska, Tanana area. One year when the steamboat was unable to continue, they ran into a caribou herd estimated to number 1 million animals, migrating across the Yukon. "They tied up for seven days waiting for the caribou to cross. They ran out of wood for the steamboats, and had to go back down 40 miles to the wood pile to pick up some more wood. On the tenth day, they came back and they said there was still caribou going across the river night and day."{{citation, title=Caribou Census Complete: 325,000 animals, date=August 2012 , url=http://westernarcticcaribou.org/wp-content/uploads/2012/08/CT2012_FINAL_0628_lowresolution.pdf , newspaper=Caribou Trails: News from the Western Arctic Caribou Herd Working Group, archive-url=https://web.archive.org/web/20120830105109/http://westernarcticcaribou.org/wp-content/uploads/2012/08/CT2012_FINAL_0628_lowresolution.pdf, location=Nome, Alaska, access-date=14 January 2014, archive-date=30 August 2012, institution=Western Arctic Caribou Herd Working Group, agency=Alaska Department of Fish and Game, url-status=live The
Gwichʼin The Gwichʼin (or Kutchin or Loucheux) are an Athabaskan languages, Athabaskan-speaking First Nations in Canada, First Nations people of Canada and an Alaskan Athabaskans, Alaska Native people. They live in the northwestern part of North America ...
, an indigenous people of northwestern Canada and northeastern Alaska, have been dependent on the international migratory
Porcupine caribou The Porcupine caribou is a herd or ecotype of the mainland barren-ground caribou (''Rangifer arcticus arcticus'', Synonym (taxonomy), syn. ''R. tarandus groenlandicus''Harding LE (2022) Available names for Rangifer (Mammalia, Artiodactyla, Cervid ...
herd for millennia.{{cite book , title=Principles of Ecosystem Stewardship: Resilience-Based Natural Resource , editor=F. Stuart Chapin III , editor2=Gary P. Kofinas , editor3=Carl Folke , year=2009 , doi=10.1007/978-0-387-73033-2 , publisher=Springer, isbn=978-0-387-73032-5 , s2cid=132900160 {{rp, 142 To them, caribou — ''vadzaih'' — is the cultural symbol and a keystone subsistence species of the Gwich'in, just as the American bison, American buffalo is to the Plains Native Americans.{{citation , title=Linguistic Team Studies Caribou Anatomy , first=Craig , last=Mishler , url=http://www.arcus.org/witness-the-arctic/2014/3/article/22797= , work=Arctic Research Consortium of the United States (ARCOS) , year=2014 , access-date=11 January 2015 , quote="A fundamental question for the research is 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." , url-status=live , archive-date=10 February 2016 , archive-url=https://web.archive.org/web/20160210131053/https://www.arcus.org/witness-the-arctic/2014/3/article/22797= Innovative Language revitalization, 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 Fairbanks, University of Alaska in Fairbanks, Alaska, 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." Elders have identified at least 150 descriptive Gwich'in names for all of the bones, organs and tissues. Associated with the caribou's anatomy are not just descriptive Gwich'in names for all of the body parts, including bones, organs, and tissues, but also "an encyclopedia of stories, songs, games, toys, ceremonies, traditional tools, skin clothing, personal names and surnames, and a highly developed ethnic cuisine." In the 1980s, Gwich'in Traditional Management Practices were established to protect the Porcupine caribou, upon which the Gwich'in depend. They "codified traditional principles of caribou management into tribal law" which include "limits on the harvest of caribou and procedures to be followed in processing and transporting caribou meat" and limits on the number of caribou to be taken per hunting trip.{{citation , first=Richard , last=Caulfield , year=1983 , title=Gwich'in Traditional Management Practices , url=http://arcticcircle.uconn.edu/ANWR/anwrgwichin1.html , series=Report to the Division of subsistence of the Alaska Department of Fish and Game , access-date=30 October 2017 , archive-url=https://web.archive.org/web/20171020081541/http://arcticcircle.uconn.edu/ANWR/anwrgwichin1.html , archive-date=20 October 2017 , url-status=live


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 people, Sámi, Nenets people, Nenets, and Komi peoples, Komi. Reindeer are used to provide renewable sources and reliable transportation. In Mongolia, the Dukha people, 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 Norway, officially the Kingdom of Norway, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. The remote Arctic island of Jan Mayen and the archipelago of Svalbard also form part of the Kingdom of ...
, 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

{{Main, Reindeer herding The reindeer is the only successfully semi-domesticated deer on a large scale in the world. Reindeer in northern Fennoscandia (northern
Norway Norway, officially the Kingdom of Norway, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. The remote Arctic island of Jan Mayen and the archipelago of Svalbard also form part of the Kingdom of ...
, Sweden and Finland) as well in the Kola Peninsula and Sakha Republic, 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 Republic, 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). DNA analysis indicates that reindeer were independently domesticated at least twice: in Fennoscandia and Western Russia (and possibly also Eastern Russia). Reindeer have been Herding, herded for centuries by several Arctic and sub-Arctic peoples, including the Sámi people, Sámi, the Nenets people, 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 Working animal, beasts of burden. Millais (1915), for example, shows a photograph (Plate LXXX) of an "Okhotsk Reindeer" saddled for riding (the rider standing behind it) beside an officer astride a steppe pony that is only slightly larger. List of domesticated animals#Tame, partially domesticated, and widely captive-bred animals, Domestic reindeer are shorter-legged and heavier than their wild counterparts.{{Citation needed, date=May 2012 In Scandinavia, management of reindeer herds is primarily conducted through ''siida'', a traditional Sámi form of cooperative association.{{Cite journal, title=Siida and traditional Sami reindeer herding knowledge , last=Korpijaakko-Mikkel, first=Sara , date=March 22, 2009 , journal=Northern Review , url=http://www.highbeam.com/doc/1G1-202252650.html, archive-url=https://web.archive.org/web/20110501035825/http://www.highbeam.com/doc/1G1-202252650.html, url-status=dead, archive-date=May 1, 2011 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 {{cvt, 20, –, 25, km a day (compared to {{cvt, 7, –, 10, km on foot, {{cvt, 70, –, 80, km by a dog sled loaded with cargo and {{cvt, 150, –, 180, km 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 {{cvt, 10, –, 14, kg 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.{{cite book , author1=United States. Bureau of Education , author2=United States. Bureau of Education. Alaska Division , title=Annual report on introduction of domestic reindeer into Alaska , url=https://books.google.com/books?id=6RcWb-PNMiMC&pg=PA18 , access-date=16 September 2011 , year=1905 , publisher=Govt. Print. Off. , pages=18–, volume=14 , url-status=live , archive-url=https://web.archive.org/web/20140101074445/http://books.google.com/books?id=6RcWb-PNMiMC&pg=PA18 , archive-date=1 January 2014 In Alaska, reindeer herders use satellite telemetry to track their herds, using online maps and databases to chart the herd's progress.{{Citation needed, date=May 2012 Domestication, 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.Europe's last wild reindeer herds in peril
{{webarchive , url=https://web.archive.org/web/20080205174121/http://www.newscientist.com/article.ns?id=dn4491 , date=5 February 2008. Newscientist. 19 December 2003. Retrieved on 16 September 2011.
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.{{cite web , url=http://reindeerherding.org/ , title=Reindeer Herding: a virtual guide to reindeer and those who herd them , institution=International Centre for Reindeer Husbandry (ICR), Government of Norway, location=Kautokeino, access-date=15 January 2014, url-status=live, archive-url=https://web.archive.org/web/20131229154038/http://reindeerherding.org/, archive-date=29 December 2013 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 ({{as of, 2001, lc=yes), the industry produces {{convert, 2,000, MT, ST 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 subsidy, 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 Photovoltaics, 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. {{blockquote, Wild reindeer "may well be the species of single greatest importance in the entire anthropological literature on hunting." Both
Aristotle Aristotle (; 384–322 BC) was an Ancient Greek philosophy, Ancient Greek philosopher and polymath. His writings cover a broad range of subjects spanning the natural sciences, philosophy, linguistics, economics, politics, psychology, a ...
and
Theophrastus Theophrastus (; ; c. 371 – c. 287 BC) was an ancient Greek Philosophy, philosopher and Natural history, naturalist. A native of Eresos in Lesbos, he was Aristotle's close colleague and successor as head of the Lyceum (classical), Lyceum, the ...
have short accounts – probably based on the same source – of an ox-sized deer species, named ''Tarand (animal), tarandos'', living in the land of the Budini, 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 The Ural Mountains ( ),; , ; , or simply the Urals, are a mountain range in Eurasia that runs north–south mostly through Russia, from the coast of the Arctic Ocean to the river Ural (river), Ural and northwestern Kazakhstan.
in c. 350 BC.{{Cite book, last1= Sarauw , first1= Georg , author-link1= Georg F.L. Sarauw , editor1-first= H. F. E. , editor1-last= Jungersen , editor2-first= E. , editor2-last= Warming , editor2-link= Eugenius Warming , title= Mindeskrift i Anledning af Hundredeaaret for Japetus Steenstrups Fødsel , chapter= Das Rentier in Europa zu den Zeiten Alexanders und Cæsars , trans-chapter= The reindeer in Europe to the times of Alexander the Great and Julius Caesar, year= 1914 , location= Copenhagen , language= de , pages= 1–33 , title-link= Japetus Steenstrup , publisher=Royal Danish Academy of Sciences and Letters 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: {{blockquote, There is an ox shaped like a Deer, stag. In the middle of its forehead a single horn grows between its ears, taller and straighter than the animal horns with which we are familiar. At the top this horn spreads out like the palm of a hand or the branches of a tree. The females are of the same form as the males, and their horns are the same shape and size. According to Olaus Magnus's ''A Description of the Northern Peoples, 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 Conrad Gessner (; ; 26 March 1516 – 13 December 1565) was a Swiss physician, naturalist, bibliographer, and philologist. Born into a poor family in Zürich, Switzerland, his father and teachers quickly realised his talents and supported him t ...
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, 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 people, Nenets, who were mobilised from the Nenets Autonomous Okrug, but reindeer herders from the Murmansk, Arkhangelsk Oblast, Arkhangelsk and Komi Republic, 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

{{Main, Santa Claus's reindeer Around the world, public interest in reindeer peaks during the Christmas season. According to Western folklore, Santa Claus's sleigh is pulled by Santa Claus's reindeer, 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:{{citation, year=2002, url=http://www.arctic.uoguelph.ca/cpl/Traditional/traditional/animals/caribou.htm, title=Tuktu — Caribou, location=Guelph, Ontario, work=Canada's Arctic, access-date=17 January 2014, url-status=live, archive-url=https://web.archive.org/web/20150923173415/http://www.arctic.uoguelph.ca/cpl/Traditional/traditional/animals/caribou.htm, archive-date=23 September 2015 {{blockquote, Once upon a time there were no caribou on the earth. But there was a man who wished for caribou, and he cut a hole deep in the ground, and up this hole came caribou, many caribou. The caribou came pouring out, until the earth was almost covered with them. And when the man thought there were caribou enough for mankind, he closed up the hole again. Thus the caribou came up on earth., 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.{{cite web, title=The ghosts on top of my head: Iconic sculpture creates campus focal point, last=Hornsby, first=Debra, date=25 August 2011, location=Banff, Alberta, access-date=31 January 2014, url=http://www.banffcentre.org/blog/2011/08/25/the-ghosts-on-top-of-my-head-iconic-sculpture-creates-campus-focal-point/, archive-url=https://web.archive.org/web/20140202204906/http://www.banffcentre.org/blog/2011/08/25/the-ghosts-on-top-of-my-head-iconic-sculpture-creates-campus-focal-point/, archive-date=2 February 2014 {{blockquote, I remember a story my Uncle Jack told me – a Dunne-Za creation story about how animals once ruled the earth and were ten times their size and that got me thinking about scale and using the idea of the antler, which is a thing that everyone is scared of, and making it into something more approachable and abstract., Brian Jungen, 2011 Tomson Highway, Order of Canada, CMTomson Highway
{{webarchive , url=https://web.archive.org/web/20110607115857/http://www.thecanadianencyclopedia.com/index.cfm?PgNm=TCE&Params=A1ARTA0010891 , date=7 June 2011. The Canadian Encyclopedia.
is a Canadians, Canadian and Cree playwright, novelist, and Children's literature, children's author, who was born in a remote area north of Brochet, Manitoba, Brochet, Manitoba. His father, Joe Highway, was a caribou hunter. His 2001 children's book entitled ''Caribou Song''/''atíhko níkamon'' was selected as one of the "Top 10 Children's Books" by the Canadian newspaper ''The Globe and Mail''. The young protagonists of ''Caribou Song'', like Tomson himself, followed the caribou herd with their families.


Heraldry and symbols

Several Norwegian municipalities have one or more reindeer depicted in their coats-of-arms: Eidfjord Municipality, Porsanger Municipality, Rendalen Municipality, Tromsø Municipality, Vadsø Municipality, and Vågå Municipality. The historic province of Västerbotten in Sweden has a reindeer in its coat of arms. The present Västerbotten County has very different borders and uses the reindeer combined with other symbols in its coat-of-arms. The city of Piteå also has a reindeer. The logo for Umeå University features three reindeer. The Canadian 25-cent coin or "Quarter (Canadian coin), 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 (St. John's), 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, Finland, Inari has a fish with reindeer antlers.Coat of arms for Inari
{{webarchive , url=https://web.archive.org/web/20130619014605/http://www.inari.fi/fi/inari-info/viestintamateriaalia/inarin-vaakuna.html , date=19 June 2013. Inari.fi.


See also

* Alaska Reindeer Service * Caribou herds and populations in Canada * Rangifer (constellation) * Rangifer (journal), ''Rangifer'' (journal) * Reindeer Police


Notes

{{notelist


References

{{Reflist, 30em


Bibliography

*{{cite web, ref=COSEWIC, title=Designatable Units for Caribou (''Rangifer tarandus'') in Canada, url=http://www.cosewic.gc.ca/eng/sct12/COSEWIC_Caribou_DU_Report_23Dec2011.pdf, website=COSEWIC, year=2011, institution=Committee on the Status of Endangered Wildlife in Canada, location=Ottawa, Ontario, access-date=18 December 2013, url-status=dead, archive-url=https://web.archive.org/web/20160303234528/http://www.cosewic.gc.ca/eng/sct12/COSEWIC_Caribou_DU_Report_23Dec2011.pdf, archive-date=3 March 2016


External links

{{Commons and category, Rangifer tarandus, Rangifer tarandus {{Wikispecies, Rangifer tarandus {{Wiktionary, reindeer, caribou * {{citation, url=http://westernarcticcaribou.org/wp-content/uploads/2012/08/CT2012_FINAL_0628_lowresolution.pdf, newspaper=Caribou Trails: News from the Western Arctic Caribou Herd Working Group, title=Caribou Census Complete: 325,000 animals, institution=Western Arctic Caribou Herd Working Group, date=August 2012, location=Nome, Alaska – the 2011 census results of the WACH, which is Alaska's largest caribou herd.
The Reindeer Portal, Source of Information About Reindeer Husbandry Worldwide





Human Role in Reindeer/Caribou Systems

Reindeer hunting as World Heritage – a ten-thousand-year-long tradition

Reindeer Research Program – Alaska reindeer research and industry development
{{Webarchive, url=https://web.archive.org/web/20210428223449/http://reindeer.salrm.uaf.edu/ , date=28 April 2021
Adaptations To Life In The Arctic
{{Webarchive, url=https://web.archive.org/web/20101212060456/http://reindeer.salrm.uaf.edu/about_reindeer/adaptations/index.php#Adaptations%20To%20Life%20In%20The%20Arctic , date=12 December 2010 – Instructional slide-show, University of Alaska
''Rangifer''
– world's only scientific journal dealing exclusively with husbandry, management and biology of Arctic and northern ungulates *{{Wikisource-inline, list= **{{Cite Americana, wstitle=Caribou, last=Ingersoll , first=Ernest , author-link=Ernest Ingersoll , short=x , noicon=x **{{Cite EB1911, wstitle=Reindeer, last=Lydekker , first=Richard , author-link=Richard Lydekker , short=x , noicon=x **{{Cite NIE, wstitle=Reindeer , year=1905 , short=x , noicon=x *{{cite web, website=United States Geological Survey, url=http://www.usgs.gov/blogs/features/usgs_top_story/the-other-364-days-of-the-year-the-real-lives-of-wild-reindeer-3/, title=The Other 364 Days of the Year: The Real Lives of Wild Reindeer Categories: Biology and Ecosystems, date=15 December 2014, author=Puckett, Catherine, author2=Landis, Ben, access-date=24 December 2014, archive-date=26 November 2015, archive-url=https://web.archive.org/web/20151126034043/http://www.usgs.gov/blogs/features/usgs_top_story/the-other-364-days-of-the-year-the-real-lives-of-wild-reindeer-3/, url-status=dead *{{cite web, website=ScienceDaily, url=https://www.sciencedaily.com/articles/r/reindeer.htm, title=Reference Article: Reindeer (caribou), access-date=25 December 2015, archive-url=https://web.archive.org/web/20150318081717/http://www.sciencedaily.com/articles/r/reindeer.htm, archive-date=18 March 2015, url-status=dead
''Growth Studies in the Reindeer'' by Charles J. Krebs
at Dartmouth College Library

University of Texas, Austin * {{cite web , title=What's the Difference: Reindeer vs. Caribou , website=National Park Service , date=October 25, 2022 , url=https://www.nps.gov/articles/000/reinderrvscaribou.htm , access-date=November 23, 2022


Caribou-specific links (North America)


Frequently Asked Questions about Caribou
from the Arctic National Wildlife Refuge
Caribou and You
– Campaign by Canadian Parks and Wilderness Society, CPAWS to protect the boreal woodland caribou, a subspecies at risk in Canada
Newfoundland Five-Year Caribou Strategy Seeks to Address Declining Populations
{{Artiodactyla, R.1 {{Working animals {{Animal domestication {{Taxonbar, from=Q39624 {{Authority control Reindeer, Alaskan cuisine Mammals described in 1758 Arctic land animals Capreolinae Mammals of the Arctic Fauna of the Holarctic realm Livestock Mammals of Asia Mammals of Canada Mammals of Greenland Mammals of Europe Mammals of Russia Mammals of the United States Pack animals Taxa named by Carl Linnaeus