The woolly mammoth (''Mammuthus primigenius'') is an extinct species of

mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus'', one of the many genera that make up the order of trunked mammals called proboscideans. The various species of mammoth were commonly equipped with long, curved tusks an ...

that lived during the

Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...

until its extinction in the

Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...

epoch. It was one of the last in a line of mammoth species, beginning with '' Mammuthus subplanifrons'' in the early

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58steppe mammoth about 800,000 years ago in East Asia. Its closest extant relative is the

Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living species of the genus ''Elephas'' and is distributed throughout the Indian subcontinent and Southeast Asia, from India in the west, Nepal in the no ...

. DNA studies show that the

Columbian mammoth The Columbian mammoth (''Mammuthus columbi'') is an extinct species of mammoth that inhabited the Americas as far north as the Northern United States and as far south as Costa Rica during the Pleistocene epoch. It was one of the last in a line ...

was a

hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...

between woolly mammoths and another lineage descended from steppe mammoths. The appearance and behaviour of this species are among the best studied of any prehistoric animal because of the discovery of frozen carcasses in

Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive region, geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a ...

and North America, as well as skeletons, teeth, stomach contents, dung, and depiction from life in prehistoric cave paintings. Mammoth remains had long been known in Asia before they became known to Europeans in the 17th century. The origin of these remains was long a matter of debate, and often explained as being remains of legendary creatures. The mammoth was identified as an extinct species of elephant by Georges Cuvier in 1796. The woolly mammoth was roughly the same size as modern African elephants. Males reached shoulder heights between and weighed up to . Females reached in shoulder heights and weighed up to . A newborn calf weighed about . The woolly mammoth was well adapted to the cold environment during the last ice age. It was covered in fur, with an outer covering of long guard hairs and a shorter undercoat. The colour of the coat varied from dark to light. The ears and tail were short to minimise

frostbite Frostbite is a skin injury that occurs when exposed to extreme low temperatures, causing the freezing of the skin or other tissues, commonly affecting the fingers, toes, nose, ears, cheeks and chin areas. Most often, frostbite occurs in the ha ...

and heat loss. It had long, curved

tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share ...

s and four

molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...

, which were replaced six times during the lifetime of an individual. Its behaviour was similar to that of modern elephants, and it used its tusks and trunk for manipulating objects, fighting, and foraging. The diet of the woolly mammoth was mainly grasses and

sedges The Cyperaceae are a family of graminoid (grass-like), monocotyledonous flowering plants known as sedges. The family is large, with some 5,500 known species described in about 90 genera, the largest being the "true sedges" genus ''Carex'' wit ...

. Individuals could probably reach the age of 60. Its habitat was the

mammoth steppe During the Last Glacial Maximum, the mammoth steppe, also known as steppe-tundra, was the Earth's most extensive biome. It spanned from Spain eastward across Eurasia to Canada and from the arctic islands southward to China. The mammoth steppe ...

, which stretched across northern Eurasia and North America. The woolly mammoth coexisted with early humans, who used its bones and tusks for making art, tools, and dwellings, and hunted the species for food. The population of woolly mammoths declined at the end of the Pleistocene, disappearing throughout most of its mainland range, although isolated populations survived on St. Paul Island until 5,600 years ago, on

Wrangel Island Wrangel Island ( rus, О́стров Вра́нгеля, r=Ostrov Vrangelya, p=ˈostrəf ˈvrangʲɪlʲə; ckt, Умӄиԓир, translit=Umqiḷir) is an island of the Chukotka Autonomous Okrug, Russia. It is the 91st largest island in the w ...

until 4,000 years ago, and possibly (based on ancient eDNA) in the

Yukon Yukon (; ; formerly called Yukon Territory and also referred to as the Yukon) is the smallest and westernmost of Canada's three territories. It also is the second-least populated province or territory in Canada, with a population of 43,964 as ...

up to 5,700 years ago and on the

Taymyr Peninsula The Taymyr Peninsula (russian: Таймырский полуостров, Taymyrsky poluostrov) is a peninsula in the Far North of Russia, in the Siberian Federal District, that forms the northernmost part of the mainland of Eurasia. Administrat ...

up to 3,900 years ago. After its extinction, humans continued using its

ivory Ivory is a hard, white material from the tusks (traditionally from elephants) and teeth of animals, that consists mainly of dentine, one of the physical structures of teeth and tusks. The chemical structure of the teeth and tusks of mammals i ...

as a raw material, a tradition that continues today. With a

genome project Genome projects are scientific endeavours that ultimately aim to determine the complete genome sequence of an organism (be it an animal, a plant, a fungus, a bacterium, an archaean, a protist or a virus) and to annotate protein-coding genes and o ...

for the mammoth completed in 2015, it has been proposed the species could be revived through various means, but none of the methods proposed are yet feasible.

Taxonomy

Remains of various extinct elephants were known by Europeans for centuries, but were generally interpreted, based on biblical accounts, as the remains of

legendary creatures A legendary creature (also mythical or mythological creature) is a type of fictional entity, typically a hybrid, that has not been proven and that is described in folklore (including myths and legends), but may be featured in historical account ...

such as behemoths or

giants A giant is a being of human appearance, sometimes of prodigious size and strength, common in folklore. Giant(s) or The Giant(s) may also refer to: Mythology and religion *Giants (Greek mythology) *Jötunn, a Germanic term often translated as 'gi ...

. They were thought to be remains of modern elephants that had been brought to Europe during the

Roman Republic The Roman Republic ( la, Res publica Romana ) was a form of government of Rome and the era of the classical Roman civilization when it was run through public representation of the Roman people. Beginning with the overthrow of the Roman Ki ...

, for example the

war elephants A war elephant was an elephant that was trained and guided by humans for combat. The war elephant's main use was to charge the enemy, break their ranks and instill terror and fear. Elephantry is a term for specific military units using elepha ...

of Hannibal and

Pyrrhus of Epirus Pyrrhus (; grc-gre, Πύρρος ; 319/318–272 BC) was a Greek king and statesman of the Hellenistic period.Plutarch. '' Parallel Lives'',Pyrrhus... He was king of the Greek tribe of Molossians, of the royal Aeacid house, and later he be ...

, or animals that had wandered north. The first woolly mammoth remains studied by European scientists were examined by

Hans Sloane Sir Hans Sloane, 1st Baronet (16 April 1660 – 11 January 1753), was an Irish physician, naturalist, and collector, with a collection of 71,000 items which he bequeathed to the British nation, thus providing the foundation of the British Mu ...

in 1728 and consisted of fossilised teeth and tusks from Siberia. Sloane was the first to recognise that the remains belonged to

elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae ...

s. Sloane turned to another biblical explanation for the presence of elephants in the Arctic, asserting that they had been buried during the

Great Flood A flood myth or a deluge myth is a myth in which a great flood, usually sent by a deity or deities, destroys civilization, often in an act of divine retribution. Parallels are often drawn between the flood waters of these myths and the primaeval ...

, and that Siberia had previously been tropical before a drastic climate change. Others interpreted Sloane's conclusion slightly differently, arguing the flood had carried elephants from the

tropics The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referr ...

to the

Arctic The Arctic ( or ) is a polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenland), Finland, Iceland, N ...

. Sloane's paper was based on travellers' descriptions and a few scattered bones collected in Siberia and Britain. He discussed the question of whether or not the remains were from elephants, but drew no conclusions. In 1738, the German zoologist

Johann Philipp Breyne Johann Philipp Breyne FRS (9 August 1680, Danzig (Gdańsk), Royal Prussia (a fief of the Crown of Poland) – 12 December 1764, Danzig, Royal Prussia), son of Jacob Breyne (1637–97), was a German-Polish botanist, palaeontologist, zoologist and ...

argued that mammoth fossils represented some kind of elephant. He could not explain why a tropical animal would be found in such a cold area as Siberia, and suggested that they might have been transported there by the Great Flood. In 1796, French biologist Georges Cuvier was the first to identify the woolly mammoth remains not as modern elephants transported to the Arctic, but as an entirely new species. He argued this species had gone extinct and no longer existed, a concept that was not widely accepted at the time. Following Cuvier's identification, German naturalist

Johann Friedrich Blumenbach Johann Friedrich Blumenbach (11 May 1752 – 22 January 1840) was a German physician, naturalist, physiologist, and anthropologist. He is considered to be a main founder of zoology and anthropology as comparative, scientific disciplines. He ...

gave the woolly mammoth its scientific name, ''Elephas primigenius'', in 1799, placing it in the same

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

as the

. This name is Latin for "the first-born elephant". Cuvier coined the name ''Elephas mammonteus'' a few months later, but the former name was subsequently used. In 1828, the British naturalist

Joshua Brookes Joshua Brookes (24 November 1761 – 10 January 1833) was a British anatomist and naturalist. Early life Brookes studied under William Hunter, William Hewson, Andrew Marshall, and John Sheldon, in London. He then attended the practice of ...

used the name ''Mammuthus borealis'' for woolly mammoth fossils in his collection that he put up for sale, thereby coining a new genus name. Mammuthus primigenius lectotypes

Where and how the word "mammoth" originated is unclear. According to the'' Oxford English Dictionary'', it comes from an old Vogul word ''mēmoŋt'', "earth-horn". It may be a version of ''mehemot'', the

Arabic Arabic (, ' ; , ' or ) is a Semitic language spoken primarily across the Arab world.Semitic languages: an international handbook / edited by Stefan Weninger; in collaboration with Geoffrey Khan, Michael P. Streck, Janet C. E.Watson; Walter ...

version of the biblical word "behemoth". Another possible origin is Estonian, where ''maa'' means "earth", and ''mutt'' means "

mole Mole (or Molé) may refer to: Animals * Mole (animal) or "true mole", mammals in the family Talpidae, found in Eurasia and North America * Golden moles, southern African mammals in the family Chrysochloridae, similar to but unrelated to Talpida ...

". The word was first used in Europe during the early 17th century, when referring to ''maimanto'' tusks discovered in Siberia. American president

Thomas Jefferson Thomas Jefferson (April 13, 1743 – July 4, 1826) was an American statesman, diplomat, lawyer, architect, philosopher, and Founding Father who served as the third president of the United States from 1801 to 1809. He was previously the natio ...

, who had a keen interest in palaeontology, was partially responsible for transforming the word "mammoth" from a noun describing the prehistoric elephant to an adjective describing anything of surprisingly large size. The first recorded use of the word as an adjective was in a description of a wheel of cheese (the " Cheshire Mammoth Cheese") given to Jefferson in 1802.Simpson, J. (2009).
Word Stories: Mammoth
." ''Oxford English Dictionary Online'', Oxford University Press. Accessed 5 June 2009. By the early 20th century, the taxonomy of extinct elephants was complex. In 1942, American palaeontologist

Henry Fairfield Osborn Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was the president of the American Museum of Natural History for 25 years and a cofounder of the American Euge ...

's posthumous monograph on the Proboscidea was published, wherein he used various taxon names that had previously been proposed for mammoth species, including replacing ''Mammuthus'' with ''Mammonteus'', as he believed the former name to be invalidly published. Mammoth taxonomy was simplified by various researchers from the 1970s onwards, all species were retained in the genus ''Mammuthus'', and many proposed differences between species were instead interpreted as intraspecific variation. Osborn chose two molars (found in Siberia and Osterode) from Blumenbach's collection at

Göttingen University Göttingen (, , ; nds, Chöttingen) is a university city in Lower Saxony, central Germany, the capital of the eponymous district. The River Leine runs through it. At the end of 2019, the population was 118,911. General information The o ...

as the

lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...

specimens for the woolly mammoth, since

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...

designation was not practised in Blumenbach's time. Soviet palaeontologist Vera Gromova further proposed the former should be considered the lectotype with the latter as paralectotype. Both molars were thought lost by the 1980s, and the more complete "Taimyr mammoth" found in Siberia in 1948 was therefore proposed as the

neotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...

specimen in 1990. Resolutions to historical issues about the validity of the genus name ''Mammuthus'' and the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...

designation of ''E. primigenius'' were also proposed. The paralectotype molar (specimen GZG.V.010.018) has since been located in the Göttingen University collection, identified by comparing it with Osborn's illustration of a cast.

Evolution

The earliest known members of the Proboscidea, the clade which contains modern elephants, existed about 55 million years ago around the

Tethys Sea The Tethys Ocean ( el, Τηθύς ''Tēthús''), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean that covered most of the Earth during much of the Mesozoic Era and early Cenozoic Era, located between the ancient continents ...

. The closest known relatives of the Proboscidea are the

sirenians The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct f ...

(dugongs and manatees) and the

hyraxes Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simi ...

(an order of small, herbivorous mammals). The family

Elephantidae Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are terrestrial large mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the ...

existed 6 million years ago in Africa and includes the modern elephants and the mammoths. Among many now extinct clades, the

mastodon A mastodon ( 'breast' + 'tooth') is any proboscidean belonging to the extinct genus ''Mammut'' (family Mammutidae). Mastodons inhabited North and Central America during the late Miocene or late Pliocene up to their extinction at the end of th ...

(''Mammut'') is only a distant relative of the mammoths, and part of the separate family

Mammutidae Mammutidae is an extinct family of proboscideans that appeared during the Oligocene epoch and survived until the start of the Holocene. The family was first described in 1922, classifying fossil specimens of the type genus ''Mammut'' ( masto ...

, which diverged 25 million years before the mammoths evolved. The following

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...

shows the placement of the genus ''Mammuthus'' among other proboscideans, based on characteristics of the hyoid bone in the neck: Mammoth molars compared

Within six weeks from 2005-2006, three teams of researchers independently assembled

mitochondrial genome Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial ...

profiles of the woolly mammoth from

ancient DNA Ancient DNA (aDNA) is DNA isolated from ancient specimens. Due to degradation processes (including cross-linking, deamination and fragmentation) ancient DNA is more degraded in comparison with contemporary genetic material. Even under the bes ...

, which allowed them to confirm the close evolutionary relationship between mammoths and Asian elephants (''Elephas maximus''). A 2015 DNA review confirmed Asian elephants as the closest living relative of the woolly mammoth. African elephants (''Loxodonta africana'') branched away from this clade around 6 million years ago, close to the time of the similar split between chimpanzees and humans. A 2010 study confirmed these relationships, and suggested the mammoth and Asian elephant lineages diverged 5.8–7.8 million years ago, while African elephants diverged from an earlier common ancestor 6.6–8.8 million years ago. In 2008, much of the woolly mammoth's

chromosomal A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...

DNA was mapped. The analysis showed that the woolly mammoth and the African elephant are 98.55% to 99.40% identical. The team mapped the woolly mammoth's nuclear genome sequence by extracting DNA from the hair follicles of both a 20,000-year-old mammoth retrieved from permafrost and another that died 60,000 years ago. In 2012, proteins were confidently identified for the first time, collected from a 43,000-year-old woolly mammoth. Since many remains of each species of mammoth are known from several localities, reconstructing the evolutionary history of the genus through morphological studies is possible. Mammoth species can be identified from the number of enamel ridges (or lamellar plates) on their

; primitive species had few ridges, and the number increased gradually as new species evolved to feed on more abrasive food items. The crowns of the teeth became deeper in height and the skulls became taller to accommodate this. At the same time, the skulls became shorter from front to back to minimise the weight of the head. The short and tall skulls of woolly and

Columbian mammoths Columbian is the adjective form of Columbia. It may refer to: Buildings * The Columbian Theatre, a music hall in northeastern Kansas * The Columbian (Chicago), a building in Illinois Published works * ''The Columbian'', a daily newspaper publis ...

(''Mammuthus columbi'') were the culmination of this process. The first known members of the genus ''Mammuthus'' are the African species '' Mammuthus subplanifrons'' from the

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58M. africanavus'' from the

. The former is thought to be the ancestor of later forms. Mammoths entered Europe around 3 million years ago. The earliest European mammoth has been named '' M. rumanus''; it spread across Europe and China. Only its molars are known, which show that it had 8–10 enamel ridges. A population evolved 12–14 ridges, splitting off from and replacing the earlier type, becoming the southern mammoth (''M. meridionalis'') about 2–1.7 million years ago. In turn, this species was replaced by the steppe mammoth (''M. trogontherii'') with 18–20 ridges, which evolved in eastern Asia around 1 million years ago. Mammoths derived from ''M. trogontherii'' evolved molars with 26 ridges 400,000 years ago in Siberia and became the woolly mammoth. Woolly mammoths entered North America about 100,000 years ago by crossing the Bering Strait.Lister, 2007. pp. 12–43

Subspecies and hybridisation

Individuals and populations showing transitional morphologies between each of the mammoth species are known, and primitive and derived species coexisted, as well, until the former disappeared. So the different species and their intermediate forms have been termed "

chronospecies A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...

". Many taxa intermediate between ''M. primigenius'' and other mammoths have been proposed, but their validity is uncertain; depending on author, they are either considered primitive forms of an advanced species or advanced forms of a primitive species. Distinguishing and determining these intermediate forms has been called one of the most long-lasting and complicated problems in Quaternary palaeontology. Regional and intermediate species and subspecies such as ''M. intermedius'', ''M. chosaricus'', ''M. p. primigenius'', ''M. p. jatzkovi'', ''M. p. sibiricus'', ''M. p. fraasi'', ''M. p. leith-adamsi'', ''M. p. hydruntinus'', ''M. p. astensis'', ''M. p. americanus'', ''M. p. compressus'' and ''M. p. alaskensis'' have been proposed. A 2011 genetic study showed that two examined specimens of the Columbian mammoth were grouped within a subclade of woolly mammoths. This suggests that the two populations interbred and produced fertile offspring. A North American type formerly referred to as ''M. jeffersonii'' may be a

between the two species. A 2015 study suggested that the animals in the range where ''M. columbi'' and ''M. primigenius'' overlapped formed a metapopulation of hybrids with varying morphology. It suggested that Eurasian ''M. primigenius'' had a similar relationship with ''M. trogontherii'' in areas where their range overlapped. In 2021, DNA older than a million years was sequenced for the first time, from two mammoth teeth of Early Pleistocene age found in eastern Siberia. One tooth from Adycha (1-1.3 million years old) belonged to a lineage that was ancestral to later woolly mammoths, whereas the other from Krestovka (1.1-1.65 million years old) belonged to new lineage, possibly a distinct species, perhaps descended from steppe mammoths that had become isolated. The study found that half of the ancestry of Columbian mammoths came from the Krestovka lineage, and the other half from woolly mammoths, with the hybridisation happening more than 420,000 years ago, during the

Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. Th ...

. Later woolly and Columbian mammoths also interbred occasionally, and mammoth species perhaps hybridised routinely when brought together by glacial expansion. These findings were the first evidence of

hybrid speciation Hybrid speciation is a form of speciation where hybridization between two different species leads to a new species, reproductively isolated from the parent species. Previously, reproductive isolation between two species and their parents was thou ...

from ancient DNA. The study also found that genetic adaptations to cold environments, such as hair growth and fat deposits, were already present in the steppe mammoth lineage, and was not unique to woolly mammoths.

Description

The appearance of the woolly mammoth is probably the best known of any prehistoric animal due to the many frozen specimens with preserved soft tissue and depictions by contemporary humans in their art. Fully grown males reached shoulder heights between and weighed up to . This is almost as large as extant male African elephants, which commonly reach a shoulder height of , and is less than the size of the earlier mammoth species ''M. meridionalis'' and ''M. trogontherii'', and the contemporary ''M. columbi''. The reason for the smaller size is unknown. Female woolly mammoths reached in shoulder heights and were built more lightly than males, weighing up to . A newborn calf would have weighed about . These sizes are deduced from comparison with modern elephants of similar size. Few frozen specimens have preserved genitals, so the sex is usually determined through examination of the skeleton. The best indication of sex is the size of the

pelvic The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The p ...

girdle, since the opening that functions as the birth canal is always wider in females than in males. Though the mammoths on Wrangel Island were smaller than those of the mainland, their size varied, and they were not small enough to be considered " island dwarfs". The last woolly mammoth populations are claimed to have decreased in size and increased their

sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...

, but this was dismissed in a 2012 study. Woolly mammoth model Royal BC Museum in Victoria

Woolly mammoths had several adaptations to the cold, most noticeably the layer of fur covering all parts of their bodies. Other adaptations to cold weather include ears that are far smaller than those of modern elephants; they were about long and across, and the ear of the 6- to 12-month-old frozen calf "Dima" was under long. The small ears reduced heat loss and

, and the tail was short for the same reason, only long in the "Berezovka mammoth". The tail contained 21 vertebrae, whereas the tails of modern elephants contain 28–33. Their skin was no thicker than that of present-day elephants, between . They had a layer of fat up to thick under the skin, which helped to keep them warm. Woolly mammoths had broad flaps of skin under their tails which covered the anus; this is also seen in modern elephants.Lister, 2007. pp. 82–87 Other characteristic features depicted in cave paintings include a large, high, single-domed head and a sloping back with a high shoulder hump; this shape resulted from the

spinous process The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...

es of the back vertebrae decreasing in length from front to rear. These features were not present in juveniles, which had convex backs like Asian elephants. Another feature shown in cave paintings was confirmed by the discovery of a frozen specimen in 1924, an adult nicknamed the "Middle Kolyma mammoth", which was preserved with a complete trunk tip. Unlike the trunk lobes of modern elephants, the upper "finger" at the tip of the trunk had a long pointed lobe and was long, while the lower "thumb" was and was broader. The trunk of "Dima" was long, whereas the trunk of the adult "Liakhov mammoth" was long. The well-preserved trunk of a juvenile specimen nicknamed "

Yuka Yuka may refer to: *Yuka (music), an Afro-Cuban style of music *Yuka (mammoth), mammoth specimen found in Yakutia, Russia *Manshu Yuka Kogyo K.K. Ssuningkai, a Japanese-German pre-WWII industrial co-operation People *Yuka (name), a Japanese perso ...

" was described in 2015, and it was shown to possess a fleshy expansion a third above the tip. Rather than oval as the rest of the trunk, this part was ellipsoidal in cross section, and double the size in diameter. The feature was shown to be present in two other specimens, of different sexes and ages.

Coat

The coat consisted of an outer layer of long, coarse "guard hair", which was on the upper part of the body, up to in length on the flanks and underside, and in diameter, and a denser inner layer of shorter, slightly curly under-wool, up to long and in diameter. The hairs on the upper leg were up to long, and those of the feet were long, reaching the toes. The hairs on the head were relatively short, but longer on the underside and the sides of the trunk. The tail was extended by coarse hairs up to long, which were thicker than the guard hairs. The woolly mammoth likely moulted seasonally, and the heaviest fur was shed during spring. Since mammoth carcasses were more likely to be preserved, possibly only the winter coat has been preserved in frozen specimens. Modern elephants have much less hair, though juveniles have a more extensive covering of hair than adults. This is thought to be for

thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...

, helping them lose heat in their hot environments. Comparison between the over-hairs of woolly mammoths and extant elephants show that they did not differ much in overall morphology. Woolly mammoths had numerous

sebaceous gland A sebaceous gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in the greatest nu ...

s in their skin, which secreted oils into their hair; this would have improved the wool's insulation, repelled water, and given the fur a glossy sheen. Preserved woolly mammoth fur is orange-brown, but this is believed to be an artefact from the bleaching of pigment during burial. The amount of

pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...

ation varied from hair to hair and within each hair. A 2006 study sequenced the ''Mc1r'' gene (which influences hair colour in mammals) from woolly mammoth bones. Two

allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chro ...

s were found: a dominant (fully active) and a

recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...

(partially active) one. In mammals, recessive'' Mc1r'' alleles result in light hair. Mammoths born with at least one copy of the dominant allele would have had dark coats, while those with two copies of the recessive allele would have had light coats. A 2011 study showed that light individuals would have been rare. A 2014 study instead indicated that the colouration of an individual varied from nonpigmented on the overhairs, bicoloured, nonpigmented and mixed red-brown guard hairs, and nonpigmented underhairs, which would give a light overall appearance.

Dentition

Woolly mammoth with downward spiralling tusk

Woolly mammoths had very long tusks (modified

incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...

teeth), which were more curved than those of modern elephants. The largest known male tusk is long and weighs , but and was a more typical size. Female tusks were smaller and thinner, and weighing . For comparison, the record for longest tusks of the African bush elephant is . The sheaths of the tusks were parallel and spaced closely. About a quarter of the length was inside the sockets. The tusks grew spirally in opposite directions from the base and continued in a curve until the tips pointed towards each other, sometimes crossing. In this way, most of the weight would have been close to the skull, and less

torque In physics and mechanics, torque is the rotational equivalent of linear force. It is also referred to as the moment of force (also abbreviated to moment). It represents the capability of a force to produce change in the rotational motion of th ...

would occur than with straight tusks. The tusks were usually asymmetrical and showed considerable variation, with some tusks curving down instead of outwards and some being shorter due to breakage. Calves developed small milk tusks a few centimetres long at six months old, which were replaced by permanent tusks a year later. Tusk growth continued throughout life, but became slower as the animal reached adulthood. The tusks grew by each year. Some cave paintings show woolly mammoths with small or no tusks, but whether this reflected reality or was artistic license is unknown. Female Asian elephants have no tusks, but no fossil evidence indicates that any adult woolly mammoths lacked them. Molaire de mammouth - font de Champdamoy

Molaire de mammouth - font de Champdamoy

Woolly mammoths had four functional molar teeth at a time, two in the upper jaw and two in the lower. About of the crown was within the jaw, and was above. The crown was continually pushed forwards and up as it wore down, comparable to a conveyor belt. The teeth had up to 26 separated ridges of enamel, which were themselves covered in "prisms" that were directed towards the chewing surface. These were quite wear-resistant and kept together by

cementum Cementum is a specialized calcified substance covering the root of a tooth. The cementum is the part of the periodontium that attaches the teeth to the alveolar bone by anchoring the periodontal ligament.Illustrated Dental Embryology, Histology, ...

and

dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by e ...

. A mammoth had six sets of molars throughout a lifetime, which were replaced five times, though a few specimens with a seventh set are known. The latter condition could extend the lifespan of the individual, unless the tooth consisted of only a few plates. The first molars were about the size of those of a human, , the third were 15 cm (6 in) long, and the sixth were about long and weighed . The molars grew larger and contained more ridges with each replacement. The woolly mammoth is considered to have had the most complex molars of any elephant.

Palaeobiology

Ice age fauna of northern Spain - Mauricio Antón

Adult woolly mammoths could effectively defend themselves from predators with their tusks, trunks and size, but juveniles and weakened adults were vulnerable to pack hunters such as

wolves The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...

, cave hyenas, and large felines. The tusks may have been used in intraspecies fighting, such as fights over

territory A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...

or mates. Display of the large tusks of males could have been used to attract females and to intimidate rivals. Because of their curvature, the tusks were unsuitable for stabbing, but may have been used for hitting, as indicated by injuries to some fossil shoulder blades. The very long hairs on the tail probably compensated for the shortness of the tail, enabling its use as a

flyswatter A fly-killing device is used for pest control of flying insects, such as houseflies, wasps, moths, gnats, and mosquitoes. Flyswatter A flyswatter (or fly-swat, fly swatter) usually consists of a small rectangular or round sheet of some acros ...

, similar to the tail on modern elephants. As in modern elephants, the sensitive and muscular trunk worked as a limb-like organ with many functions. It was used for manipulating objects, and in social interactions. The well-preserved foot of the adult male " Yukagir mammoth" shows that the soles of the feet contained many cracks that would have helped in gripping surfaces during locomotion. Like modern elephants, woolly mammoths walked on their toes and had large, fleshy pads behind the toes. Like modern elephants, woolly mammoths were likely very social and lived in matriarchal (female-led) family groups. This is supported by fossil assemblages and cave paintings showing groups, implying that most of their other social behaviours were likely similar to those of modern elephants. How many mammoths lived at one location at a time is unknown, as fossil deposits are often accumulations of individuals that died over long periods of time. The numbers likely varied by season and lifecycle events. Modern elephants can form large herds, sometimes consisting of multiple family groups, and these herds can include thousands of animals migrating together. Mammoths may have formed large herds more often, since animals that live in open areas are more likely to do this than those in forested areas. Trackways made by a woolly mammoth herd 11,300–11,000 years ago have been found in the St. Mary Reservoir in Canada, showing that in this case almost equal numbers of adults, subadults, and juveniles were found. The adults had a stride of , and the juveniles ran to keep up.

Adaptations to cold

The woolly mammoth was probably the most specialised member of the family Elephantidae. In addition to their fur, they had lipopexia (fat storage) in their neck and

withers The withers is the ridge between the shoulder blades of an animal, typically a quadruped. In many species, it is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, cattle a ...

, for times when food availability was insufficient during winter, and their first three molars grew more quickly than in the calves of modern elephants. The expansion identified on the trunk of "Yuka" and other specimens was suggested to function as a "fur mitten"; the trunk tip was not covered in fur, but was used for foraging during winter, and could have been heated by curling it into the expansion. The expansion could be used to melt snow if a shortage of water to drink existed, as melting it directly inside the mouth could disturb the thermal balance of the animal. As in

reindeer Reindeer (in North American English, known as caribou if wild and ''reindeer'' if domesticated) are deer in the genus ''Rangifer''. For the last few decades, reindeer were assigned to one species, ''Rangifer tarandus'', with about 10 sub ...

and

musk oxen Musk ( Persian: مشک, ''Mushk'') is a class of aromatic substances commonly used as base notes in perfumery. They include glandular secretions from animals such as the musk deer, numerous plants emitting similar fragrances, and artificial sub ...

, the

haemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...

of the woolly mammoth was adapted to the cold, with three mutations to improve oxygen delivery around the body and prevent freezing. This feature may have helped the mammoths to live at high latitudes. In a 2015 study, high-quality

genome In the fields of molecular biology and genetics, 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 g ...

sequences from three Asian elephants and two woolly mammoths were compared. About 1.4 million DNA nucleotide differences were found between mammoths and elephants, which affect the sequence of more than 1,600 proteins. Differences were noted in genes for a number of aspects of physiology and biology that would be relevant to Arctic survival, including development of skin and hair, storage and metabolism of adipose tissue, and perceiving temperature. Genes related to both sensing temperature and transmitting that sensation to the brain were altered. One of the heat-sensing genes encodes a protein, TRPV3, found in skin, which affects hair growth. When inserted into human cells, the mammoth's version of the protein was found to be less sensitive to heat than the elephant's. This is consistent with a previous observation that mice lacking active TRPV3 are likely to spend more time in cooler cage locations than wild-type mice, and have wavier hair. Several alterations in circadian clock genes were found, perhaps needed to cope with the extreme polar variation in length of daylight. Similar mutations are known in other Arctic mammals, such as reindeer. A 2019 study of the woolly mammoth mitogenome suggest that these had metabolic adaptations related to extreme environments.

Diet

Food at various stages of digestion has been found in the intestines of several woolly mammoths, giving a good picture of their diet. Woolly mammoths sustained themselves on plant food, mainly grasses and sedges, which were supplemented with

herbaceous plant Herbaceous plants are vascular plants that have no persistent wood, woody stems above ground. This broad category of plants includes many perennial plant, perennials, and nearly all Annual plant, annuals and Biennial plant, biennials. Definition ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s, shrubs, mosses, and tree matter. The composition and exact varieties differed from location to location. Woolly mammoths needed a varied diet to support their growth, like modern elephants. An adult of 6 tons would need to eat daily, and may have foraged as long as 20 hours every day. The two-fingered tip of the trunk was probably adapted for picking up the short grasses of the last

ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...

(

Quaternary glaciation The Quaternary glaciation, also known as the Pleistocene glaciation, is an alternating series of glacial and interglacial periods during the Quaternary period that began 2.58 Ma (million years ago) and is ongoing. Although geologists describ ...

, 2.58 million years ago to present) by wrapping around them, whereas modern elephants curl their trunks around the longer grass of their tropical environments. The trunk could be used for pulling off large grass tufts, delicately picking

bud In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be spec ...

s and flowers, and tearing off leaves and branches where trees and shrubs were present. The "Yukagir mammoth" had ingested plant matter that contained spores of dung fungus. Isotope analysis shows that woolly mammoths fed mainly on C3 plants, unlike horses and rhinos. Scientists identified milk in the stomach and faecal matter in the intestines of the mammoth calf " Lyuba". The faecal matter may have been eaten by "Lyuba" to promote development of the intestinal microbes necessary for digestion of vegetation, as is the case in modern elephants. An isotope analysis of woolly mammoths from Yukon showed that the young nursed for at least 3 years, and were

weaned Weaning is the process of gradually introducing an infant human or another mammal to what will be its adult diet while withdrawing the supply of its mother's milk. The process takes place only in mammals, as only mammals produce milk. The infan ...

and gradually changed to a diet of plants when they were 2–3 years old. This is later than in modern elephants and may be due to a higher risk of predator attack or difficulty in obtaining food during the long periods of winter darkness at high latitudes. Worn Woolly mammoth tusk

The molars were adapted to their diet of coarse tundra grasses, with more enamel plates and a higher crown than their earlier, southern relatives. The woolly mammoth chewed its food by using its powerful jaw muscles to move the mandible forwards and close the mouth, then backwards while opening; the sharp enamel ridges thereby cut across each other, grinding the food. The ridges were wear-resistant to enable the animal to chew large quantities of food, which often contained grit. Woolly mammoths may have used their tusks as shovels to clear snow from the ground and reach the vegetation buried below, and to break ice to drink. This is indicated on many preserved tusks by flat, polished sections up to long, as well as scratches, on the part of the surface that would have reached the ground (especially at their outer curvature). The tusks were used for obtaining food in other ways, such as digging up plants and stripping off bark.

Life history

Cross sections of African elephant and woolly mammoth tusks

The lifespan of mammals is related to their size, and since modern elephants can reach the age of 60 years, the same is thought to be true for woolly mammoths, which were of a similar size. The age of a mammoth can be roughly determined by counting the

growth rings Dendrochronology (or tree-ring dating) is the scientific method of dating tree rings (also called growth rings) to the exact year they were formed. As well as dating them, this can give data for dendroclimatology, the study of climate and atmos ...

of its tusks when viewed in cross section, but this does not account for its early years, as these are represented by the tips of the tusks, which are usually worn away. In the remaining part of the tusk, each major line represents a year, and weekly and daily ones can be found in between. Dark bands correspond to summers, so determining the season in which a mammoth died is possible. The growth of the tusks slowed when foraging became harder, for example during winter, during disease, or when a male was banished from the herd (male elephants live with their herds until about the age of 10). Mammoth tusks dating to the harshest period of the last glaciation 25–20,000 years ago show slower growth rates.Lister, 2007. pp. 83–107. Woolly mammoths continued growing past adulthood, like other elephants. Unfused limb bones show that males grew until they reached the age of 40, and females grew until they were 25. The frozen calf "Dima" was tall when it died at the age of 6–12 months. At this age, the second set of molars would be in the process of erupting, and the first set would be worn out at 18 months of age. The third set of molars lasted for 10 years, and this process was repeated until the final, sixth set emerged when the animal was 30 years old. When the last set of molars was worn out, the animal would be unable to chew and feed, and it would die of starvation. A study of North American mammoths found that they often died during winter or spring, the hardest times for northern animals to survive. Examination of preserved calves shows that they were all born during spring and summer, and since modern elephants have gestation periods of 21–22 months, the

mating season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and ch ...

probably was from summer to autumn. δ15N isotopic analysis of the teeth of "Lyuba" has demonstrated their prenatal development, and indicates its gestation period was similar to that of a modern elephant, and that it was born in spring. The best-preserved head of a frozen adult specimen, that of a male nicknamed the "Yukagir mammoth", shows that woolly mammoths had temporal glands between the ear and the eye. This feature indicates that, like bull elephants, male woolly mammoths entered "

musth Musth or must (from Persian, )''The Oxford Dictionary and Thesaurus: American edition'', published 1996 by Oxford University Press; p. 984 is a periodic condition in bull (male) elephants characterized by aggressive behavior and accompanied ...

", a period of heightened aggressiveness. The glands are used especially by males to produce an oily substance with a strong smell called

temporin Temporins are a family of peptides isolated originally from the skin secretion of the European red frog, ''Rana temporaria''. Peptides belonging to the temporin family have been isolated also from closely related North American frogs, such as ''R ...

. Their fur may have helped in spreading the scent further.

Palaeopathology

Evidence of several different

bone disease Bone disease refers to the medical conditions which affect the bone. Terminology A bone disease is also called an "osteopathy", but because the term osteopathy is often used to refer to an alternative health-care philosophy, use of the term can ...

s has been found in woolly mammoths. The most common of these was

osteoarthritis Osteoarthritis (OA) is a type of degenerative joint disease that results from breakdown of joint cartilage and underlying bone which affects 1 in 7 adults in the United States. It is believed to be the fourth leading cause of disability in the w ...

, found in 2% of specimens. One specimen from Switzerland had several fused

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...

e as a result of this condition. The "Yukagir mammoth" had suffered from

spondylitis Spondylitis is an inflammation of the vertebrae. It is a form of spondylopathy. In many cases, spondylitis involves one or more vertebral joints, as well, which itself is called spondylarthritis. __TOC__ Types Pott disease is a tuberculous dis ...

in two vertebrae, and

osteomyelitis Osteomyelitis (OM) is an infection of bone. Symptoms may include pain in a specific bone with overlying redness, fever, and weakness. The long bones of the arms and legs are most commonly involved in children e.g. the femur and humerus, while the ...

is known from some specimens. Several specimens have healed bone fractures, showing that the animals had survived these injuries. An abnormal number of cervical vertebrae has been found in 33% of specimens from the North Sea region, probably due to inbreeding in a declining population.

Parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...

flies and protozoa were identified in the gut of the calf "Dima". Distortion in the molars is the most common health problem found in woolly mammoth fossils. Sometimes, the replacement was disrupted, and the molars were pushed into abnormal positions, but some animals are known to have survived this. Teeth from Britain showed that 2% of specimens had

periodontal disease Periodontal disease, also known as gum disease, is a set of inflammatory conditions affecting the tissues surrounding the teeth. In its early stage, called gingivitis, the gums become swollen and red and may bleed. It is considered the main caus ...

, with half of these containing

caries Tooth decay, also known as cavities or caries, is the breakdown of teeth due to acids produced by bacteria. The cavities may be a number of different colors from yellow to black. Symptoms may include pain and difficulty with eating. Complicatio ...

. The teeth sometimes had

cancerous Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal bl ...

growths.

Distribution and habitat

The habitat of the woolly mammoth is known as "

" or "tundra steppe". This environment stretched across northern Asia, many parts of Europe, and the northern part of North America during the last ice age. It was similar to the grassy

steppes In physical geography, a steppe () is an ecoregion characterized by grassland plains without trees apart from those near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the temperate grassland ...

of modern Russia, but the

flora Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous (ecology), indigenous) native plant, native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' ...

was more diverse, abundant, and grew faster. Grasses, sedges, shrubs, and herbaceous plants were present, and scattered trees were mainly found in southern regions. This habitat was not dominated by ice and snow, as is popularly believed, since these regions are thought to have been

high-pressure area A high-pressure area, high, or anticyclone, is an area near the surface of a planet where the atmospheric pressure is greater than the pressure in the surrounding regions. Highs are middle-scale meteorological features that result from interpl ...

s at the time. The habitat of the woolly mammoth supported other grazing herbivores such as the

woolly rhinoceros The woolly rhinoceros (''Coelodonta antiquitatis'') is an extinct species of rhinoceros that was common throughout Europe and Asia during the Pleistocene epoch and survived until the end of the last glacial period. The woolly rhinoceros was a me ...

, wild horses, and bison.Lister, 2007. pp. 88–89 The Altai-Sayan assemblages are the modern biomes most similar to the "mammoth steppe". A 2014 study concluded that

forb A forb or phorb is an herbaceous flowering plant that is not a graminoid (grass, sedge, or rush). The term is used in biology and in vegetation ecology, especially in relation to grasslands and understory. Typically these are dicots without woo ...

s (a group of herbaceous plants) were more important in the steppe-tundra than previously acknowledged, and that it was a primary food source for the ice-age megafauna. Wooly Mammoths

The southernmost woolly mammoth specimen known is from the Shandong province of China, and is 33,000 years old. The southernmost European remains are from the Depression of Granada in Spain and are of roughly the same age. DNA studies have helped determine the

phylogeography Phylogeography is the study of the historical processes that may be responsible for the past to present geographic distributions of genealogical lineages. This is accomplished by considering the geographic distribution of individuals in light of ge ...

of the woolly mammoth. A 2008 DNA study showed two distinct groups of woolly mammoths: one that became extinct 45,000 years ago and another one that became extinct 12,000 years ago. The two groups are speculated to be divergent enough to be characterised as subspecies. The group that became extinct earlier stayed in the middle of the high Arctic, while the group with the later extinction had a much wider range. Recent stable isotope studies of Siberian and New World mammoths have shown there were differences in climatic conditions on either side of the

Bering land bridge 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 72 degrees north latitude in the Chukchi Sea; and on the south by the tip of ...

(Beringia), with Siberia being more uniformly cold and dry throughout the Late Pleistocene. During the

Younger Dryas The Younger Dryas (c. 12,900 to 11,700 years BP) was a return to glacial conditions which temporarily reversed the gradual climatic warming after the Last Glacial Maximum (LGM, c. 27,000 to 20,000 years BP). The Younger Dryas was the last stag ...

age, woolly mammoths briefly expanded into north-east Europe, whereafter the mainland populations became extinct. A 2008 genetic study showed that some of the woolly mammoths that entered North America through the Bering land bridge from Asia migrated back about 300,000 years ago and had replaced the previous Asian population by about 40,000 years ago, not long before the entire species became extinct. Fossils of woolly mammoths and Columbian mammoths have been found together in a few localities of North America, including the Hot Springs sinkhole of

South Dakota South Dakota (; Sioux: , ) is a U.S. state in the North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Lakota and Dakota Sioux Native American tribes, who comprise a large porti ...

where their regions overlapped. It is unknown whether the two species were sympatric and lived there simultaneously, or if the woolly mammoths may have entered these southern areas during times when Columbian mammoth populations were absent there.

Relationship with humans

Modern humans co-existed with woolly mammoths during the

Upper Palaeolithic The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories coin ...

period when the humans entered Europe from Africa between 30,000 and 40,000 years ago. Before this, Neanderthals had co-existed with mammoths during the

Middle Palaeolithic The Middle Paleolithic (or Middle Palaeolithic) is the second subdivision of the Paleolithic or Old Stone Age as it is understood in Europe, Africa and Asia. The term Middle Stone Age is used as an equivalent or a synonym for the Middle Pale ...

and already used mammoth bones for tool-making and building materials. Woolly mammoths were very important to ice age humans, and human survival may have depended on the mammoth in some areas. Evidence for such co-existence was not recognised until the 19th century.

William Buckland William Buckland DD, FRS (12 March 1784 – 14 August 1856) was an English theologian who became Dean of Westminster. He was also a geologist and palaeontologist. Buckland wrote the first full account of a fossil dinosaur, which he named ' ...

published his discovery of the

Red Lady of Paviland The Red Lady of Paviland is an Upper Paleolithic partial skeleton of a male dyed in red ochre and buried in Wales 33,000 BP. The bones were discovered in 1823 by William Buckland in an archaeological dig at Goat's Hole Cave (Paviland cave) – ...

skeleton in 1823, which was found in a cave alongside woolly mammoth bones, but he mistakenly denied that these were contemporaries. In 1864,

Édouard Lartet Édouard Lartet (15 April 180128 January 1871) was a French geologist and paleontologist, and a pioneer of Paleolithic archaeology. Biography Lartet was born near Castelnau-Barbarens, ' of Gers, France, where his family had lived for more than ...

found an engraving of a woolly mammoth on a piece of mammoth ivory in the

Abri de la Madeleine The archaeological site Abri de la Madeleine (Magdalene Shelter) is a rock shelter under an overhanging cliff situated near Tursac, in the Dordogne ''département'' of the Aquitaine Région of South-Western France. It represents the type site of ...

cave in

Dordogne Dordogne ( , or ; ; oc, Dordonha ) is a large rural department in Southwestern France, with its prefecture in Périgueux. Located in the Nouvelle-Aquitaine region roughly half-way between the Loire Valley and the Pyrenees, it is name ...

, France. The engraving was the first widely accepted evidence for the co-existence of humans with prehistoric extinct animals and is the first contemporary depiction of such a creature known to modern science. The woolly mammoth is the third-most depicted animal in ice age art, after horses and bison, and these images were produced between 35,000 and 11,500 years ago. Today, more than 500 depictions of woolly mammoths are known, in media ranging from cave paintings and engravings on the walls of 46 caves in Russia, France, and Spain to engravings and sculptures (termed " portable art") made from ivory, antler, stone and bone. Cave paintings of woolly mammoths exist in several styles and sizes. The French

Rouffignac Cave The Rouffignac cave, in the French commune of Rouffignac-Saint-Cernin-de-Reilhac in the Dordogne département, contains over 250 engravings and cave paintings dating back to the Upper Paleolithic. In conjunction with other caves and abris of th ...

has the most depictions, 159, and some of the drawings are more than in length. Other notable caves with mammoth depictions are the

Chauvet Cave The Chauvet-Pont-d'Arc Cave (french: Grotte Chauvet-Pont d'Arc, ) in the Ardèche department of southeastern France is a cave that contains some of the best-preserved figurative cave paintings in the world, as well as other evidence of Upper Pale ...

, Les Combarelles Cave, and

Font-de-Gaume Font-de-Gaume is a cave near Les Eyzies-de-Tayac-Sireuil in the Dordogne départment of south-west France. The cave contains prehistoric polychrome cave paintings and engravings dating to the Magdalenian period. Discovered in 1901, more th ...

.Lister, 2007. pp. 118–125 A depiction in the

Cave of El Castillo The Cueva del Castillo, or Cave of the Castle, is an archaeological site within the complex of the Caves of Monte Castillo, in Puente Viesgo, Cantabria, Spain. The archaeological stratigraphy has been divided into around 19 layers, dependin ...

may instead show ''Palaeoloxodon'', the "straight-tusked elephant". "Portable art" can be more accurately dated than cave art since it is found in the same deposits as tools and other ice age artefacts. The largest collection of portable mammoth art, consisting of 62 depictions on 47 plaques, was found in the 1960s at an excavated open-air camp near Gönnersdorf in Germany. A correlation between the number of mammoths depicted and the species that were most often hunted does not seem to exist, since reindeer bones are the most frequently found animal remains at the site. Two

spear throwers A spear-thrower, spear-throwing lever or ''atlatl'' (pronounced or ; Classical Nahuatl, Nahuatl ''ahtlatl'' ) is a tool that uses leverage to achieve greater velocity in Dart (missile), dart or javelin-throwing, and includes a Plain bearing, b ...

shaped as woolly mammoths have been found in France. Some portable mammoth depictions may not have been produced where they were discovered, but could have moved around by ancient trading.

Exploitation

Woolly mammoth bones were used as construction material for dwellings by both Neanderthals and modern humans during the ice age. More than 70 such dwellings are known, mainly from the

East European Plain The East European Plain (also called the Russian Plain, "Extending from eastern Poland through the entire European Russia to the Ural Mountaina, the ''East European Plain'' encompasses all of the Baltic states and Belarus, nearly all of Ukraine, an ...

. The bases of the huts were circular, and ranged from . The arrangement of dwellings varied, and ranged from apart, depending on location. Large bones were used as foundations for the huts, tusks for the entrances, and the roofs were probably skins held in place by bones or tusks. Some huts had floors that extended below ground. Some of the bones used for materials may have come from mammoths killed by humans, but the state of the bones, and the fact that bones used to build a single dwelling varied by several thousands of years in age, suggests that they were collected remains of long-dead animals. Woolly mammoth bones were made into various tools, furniture, and musical instruments. Large bones, such as shoulder blades, were used to cover dead human bodies during burial. Woolly mammoth ivory was used to create art objects. Several

Venus figurines A Venus figurine is any Upper Palaeolithic statuette portraying a woman, usually carved in the round.Fagan, Brian M., Beck, Charlotte, "Venus Figurines", ''The Oxford Companion to Archaeology'', 1996, Oxford University Press, pp. 740–741 Most ...

, including the

Venus of Brassempouy The Venus of Brassempouy (French: ''la Dame de Brassempouy'', , meaning "Lady of Brassempouy", or ''Dame à la Capuche'', "Lady with the Hood") is a fragmentary ivory figurine from the Upper Palaeolithic, apparently broken from a larger figure at ...

and the

Venus of Lespugue The Venus of Lespugue is a Venus figurine, a statuette of a nude female figure of the Gravettian, dated to between 26,000 and 24,000 years ago. Discovery It was discovered in 1922 in the Rideaux cave of Lespugue (Haute-Garonne) in the foothills ...

, were made from this material. Weapons made from ivory, such as daggers, spears, and a

boomerang A boomerang () is a thrown tool, typically constructed with aerofoil sections and designed to spin about an axis perpendicular to the direction of its flight. A returning boomerang is designed to return to the thrower, while a non-returning ...

, are known. A 2019 study found that woolly mammoth ivory was the most suitable bony material for the production of big game projectile points during the Late Plesistocene. To be able to process the ivory, the large tusks had to be chopped, chiseled, and split into smaller, more manageable pieces. Some ivory artefacts show that tusks had been straightened, and how this was achieved is unknown. Several woolly mammoth specimens show evidence of being butchered by humans, which is indicated by breaks, cut marks, and associated stone tools. How much prehistoric humans relied on woolly mammoth meat is unknown, since many other large herbivores were available. Many mammoth carcasses may have been scavenged by humans rather than hunted. Some cave paintings show woolly mammoths in structures interpreted as pitfall traps. Few specimens show direct, unambiguous evidence of having been hunted by humans. A Siberian specimen with a spearhead embedded in its shoulder blade shows that a spear had been thrown at it with great force. At a site in southern Polan that contains bones from over 100 mammoths, stone spear tips have been found embedded in bones, and many stone spear points in the site were damaged from impact against mammoth bones, indicating that mammoths were the major prey for people at the time. A specimen from the

Mousterian The Mousterian (or Mode III) is an archaeological industry of stone tools, associated primarily with the Neanderthals in Europe, and to the earliest anatomically modern humans in North Africa and West Asia. The Mousterian largely defines the l ...

age of Italy shows evidence of spear hunting by Neanderthals. The juvenile specimen nicknamed "Yuka" is the first frozen mammoth with evidence of human interaction. It shows evidence of having been killed by a large predator, and of having been scavenged by humans shortly after. Some of its bones had been removed, and were found nearby. A site near the

Yana River The Yana ( rus, Я́на, p=ˈjanə; sah, Дьааҥы, ''Caañı'') is a river in Sakha in Russia, located between the Lena to the west and the Indigirka to the east. Course It is long, and its drainage basin covers . Including its longes ...

in Siberia has revealed several specimens with evidence of human hunting, but the finds were interpreted to show that the animals were not hunted intensively, but perhaps mainly when ivory was needed. Two woolly mammoths from

Wisconsin Wisconsin () is a state in the upper Midwestern United States. Wisconsin is the 25th-largest state by total area and the 20th-most populous. It is bordered by Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake M ...

, the "Schaefer" and "Hebior mammoths", show evidence of having been butchered by Palaeoamericans.

Extinction

Projectile points made from mammoth ivory

Most woolly mammoth populations disappeared during the late Pleistocene and mid-

, alongside most of the

Pleistocene megafauna Pleistocene megafauna is the set of large animals that lived on Earth during the Pleistocene epoch. Pleistocene megafauna became extinct during the Quaternary extinction event resulting in substantial changes to ecosystems globally. The role of ...

(including the Columbian mammoth). This extinction formed part of the

Quaternary extinction event The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecolog ...

, which began 40,000 years ago and peaked between 14,000 and 11,500 years ago. Scientists are divided over whether hunting or climate change, which led to the shrinkage of its habitat, was the main factor that contributed to the extinction of the woolly mammoth, or whether it was due to a combination of the two. Whatever the cause, large mammals are generally more vulnerable than smaller ones due to their smaller population size and low reproduction rates. Different woolly mammoth populations did not die out simultaneously across their range, but gradually became extinct over time. Most populations disappeared between 14,000 and 10,000 years ago. The youngest fossils of the mainland population are from the Kyttyk Peninsula of Siberia and date to 9,650 years ago. A small population of woolly mammoths survived on St. Paul Island, Alaska, well into the Holocene with the most recently published date of extinction being 5,600 years B.P. The last population known from fossils remained on

in the

Arctic Ocean The Arctic Ocean is the smallest and shallowest of the world's five major oceans. It spans an area of approximately and is known as the coldest of all the oceans. The International Hydrographic Organization (IHO) recognizes it as an ocean, a ...

until 4,000 years ago, well into the start of human

civilization A civilization (or civilisation) is any complex society characterized by the development of a state, social stratification, urbanization, and symbolic systems of communication beyond natural spoken language (namely, a writing system). ...

and concurrent with the construction of the

Great Pyramid The Great Pyramid of Giza is the biggest Egyptian pyramid and the tomb of Fourth Dynasty pharaoh Khufu. Built in the early 26th century BC during a period of around 27 years, the pyramid is the oldest of the Seven Wonders of the Ancient World ...

of ancient Egypt. However, ancient genetic evidence supports the existence of small mainland populations that died out at around the same time as their island counterparts; two studies in 2021 found that based on eDNA, mammoths survived in the

until about 5,700 years ago, roughly concurrent with the St. Paul population, and on the

of Siberia until 3,900 to 4,100 years ago, roughly concurrent with the Wrangel population. The Taymyr Peninsula, with its drier habitat, may have served as a refugium for the mammoth steppe, supporting mammoths and other widespread Ice Age mammals such as wild horses ('' Equus'' sp.). DNA sequencing of remains of two mammoths, one from Siberia 44,800 years BP and one from Wrangel Island 4,300 years BP, indicates two major population crashes: one around 280,000 years ago from which the population recovered, and a second about 12,000 years ago, near the ice age's end, from which it did not. The Wrangel Island mammoths were isolated for 5000 years by rising post-ice-age sea level, and resultant inbreeding in their small population of about 300 to 1000 individuals led to a 20% to 30% loss of heterozygosity, and a 65% loss in mitochondrial DNA diversity. The population seems to have subsequently been stable, without suffering further significant loss of genetic diversity. Genetic evidence thus implies the extinction of this final population was sudden, rather than the culmination of a gradual decline. Woolly Mammoth Climatic Suitability - Nogués-Bravo 2008

Woolly Mammoth Climatic Suitability - Nogués-Bravo 2008

Before their extinction, the Wrangel Island mammoths had accumulated numerous genetic defects due to their small population; in particular, a number of genes for olfactory receptors and urinary proteins became nonfunctional, possibly because they had lost their selective value on the island environment. It is not clear whether these genetic changes contributed to their extinction. It has been proposed that these changes are consistent with the concept of genomic meltdown; however, the sudden disappearance of an apparently stable population may be more consistent with a catastrophic event, possibly related to climate (such as icing of the snowpack) or a human hunting expedition. The disappearance coincides roughly in time with the first evidence for humans on the island. The woolly mammoths of eastern Beringia (modern

Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S. ...

and

) had similarly died out about 13,300 years ago, soon (roughly 1000 years) after the first appearance of humans in the area, which parallels the fate of all the other late Pleistocene proboscids (mammoths, gomphotheres, and mastodons), as well as most of the rest of the megafauna, of the

Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...

. In contrast, the St. Paul Island mammoth population apparently died out before human arrival because of habitat shrinkage resulting from the post-ice age sea-level rise, perhaps in large measure as a result of a consequent reduction in the freshwater supply. Changes in climate shrank suitable mammoth habitat from 42,000 years ago to 6,000 years ago. Woolly mammoths survived an even greater loss of habitat at the end of the

Saale glaciation The Saale glaciation or Saale Glaciation, sometimes referred to as the Saalian glaciation, Saale cold period (german: Saale-Kaltzeit), Saale complex (''Saale-Komplex'') or Saale glacial stage (''Saale-Glazial'', colloquially also the ''Saale-Eiszei ...

125,000 years ago, and humans likely hunted the remaining populations to extinction at the end of the last glacial period. Studies of an 11,300–11,000-year-old

trackway Historic roads (historic trails in USA and Canada) are paths or routes that have historical importance due to their use over a period of time. Examples exist from prehistoric times until the early 20th century. They include ancient trackways ...

in south-western Canada showed that ''M. primigenius'' was in decline while coexisting with humans, since far fewer tracks of juveniles were identified than would be expected in a normal herd. A 2021 study indicates, however, that although humans likely exerted a significant selective pressure on mammoths that led to them going extinct earlier than they otherwise would have, the final impetus for mammoth extinction was likely vegetation changes caused by a changed precipitation regime at the end of the Ice Age. The decline of the woolly mammoth could have increased temperatures by up to at high latitudes in the Northern Hemisphere. Mammoths frequently ate birch trees, creating a grassland habitat. With the disappearance of mammoths, birch forests, which absorb more sunlight than grasslands, expanded, leading to regional warming.

Fossil specimens

Woolly mammoth fossils have been found in many different types of deposits, including former rivers and lakes, and in "

Doggerland Doggerland was an area of land, now submerged beneath the North Sea, that connected Britain to continental Europe. It was flooded by rising sea levels around 6500–6200 BCE. The flooded land is known as the Dogger Littoral. Geological sur ...

" in the

North Sea The North Sea lies between Great Britain, Norway, Denmark, Germany, the Netherlands and Belgium. An epeiric sea, epeiric sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the ...

, which was dry at times during the ice age. Such fossils are usually fragmentary and contain no soft tissue. Accumulations of modern elephant remains have been termed "

elephants' graveyard An elephants' graveyard (also written elephant graveyard, elephant's graveyard, or elephants' cemetery) is a place where, according to legend, older elephants instinctively direct themselves when they reach a certain age. According to this legend, ...

s", as these sites were erroneously thought to be where old elephants went to die. Similar accumulations of woolly mammoth bones have been found; these are thought to be the result of individuals dying near or in the rivers over thousands of years, and their bones eventually being brought together by the streams. Some accumulations are thought to be the remains of herds that died together at the same time, perhaps due to flooding. Natural traps, such as kettle holes, sink holes, and mud, have trapped mammoths in separate events over time. 200805231215 Wollhaarmammut Millie Schädel

200805231215 Wollhaarmammut Millie Schädel

Apart from frozen remains, the only soft tissue known is from a specimen that was preserved in a

petroleum seep A petroleum seep is a place where natural liquid or gaseous hydrocarbons escape to the earth's atmosphere and surface, normally under low pressure or flow. Seeps generally occur above either terrestrial or offshore petroleum accumulation stru ...

in Starunia, Poland. Frozen remains of woolly mammoths have been found in the northern parts of

and Alaska, with far fewer finds in the latter. Such remains are mostly found above the Arctic Circle, in permafrost. Soft tissue apparently was less likely to be preserved between 30,000 and 15,000 years ago, perhaps because the climate was milder during that period. Most specimens have partially degraded before discovery, due to exposure or to being scavenged. This " natural mummification" required the animal to have been buried rapidly in liquid or semisolids such as silt, mud, and icy water, which then froze.Lister, 2007. pp. 50–53 The presence of undigested food in the stomach and seed pods still in the mouth of many of the specimens suggests neither starvation nor exposure is likely. The maturity of this ingested vegetation places the time of death in autumn rather than in spring, when flowers would be expected. The animals may have fallen through ice into small ponds or potholes, entombing them. Many are certainly known to have been killed in rivers, perhaps through being swept away by floods. In one location, by the Byoryolyokh River in

Yakutia Sakha, officially the Republic of Sakha (Yakutia),, is the largest republic of Russia, located in the Russian Far East, along the Arctic Ocean, with a population of roughly 1 million. Sakha comprises half of the area of its governing Far E ...

in Siberia, more than 8,000 bones from at least 140 mammoths have been found in a single spot, apparently having been swept there by the current.

Frozen specimens

Between 1692 and 1806, a handful of reports of frozen mammoth remains with soft tissue were published reached Europe, though none were collected during that time. While frozen woolly mammoth carcasses had been excavated by Europeans as early as 1728, the first fully documented specimen was discovered near the delta of the Lena River in 1799 by Ossip Schumachov, a Siberian hunter. While in Yakutsk in 1806,

Michael Friedrich Adams Johann Friedrich Adam, later called Michael Friedrich Adams (1780 in Moscow – 1 March 1838, in Vereya) was a botanist from Saint Petersburg, St. Petersburg, Russia. He studied from 1795–1796 in the medical school of Saint Petersburg, S ...

heard about the frozen mammoth. Adams recovered the entire skeleton, apart from the tusks, which Shumachov had already sold, and one foreleg, most of the skin, and nearly 18 kg (40 lb) of hair. During his return voyage, he purchased a pair of tusks that he believed were the ones that Shumachov had sold. Adams brought all to the

Zoological Museum of the Zoological Institute of the Russian Academy of Sciences The Zoological Museum of the Zoological Institute of the Russian Academy of Sciences is a Russian museum devoted to zoology. It is located in Saint Petersburg, on Universitetskaya Embankment. It is one of the ten largest nature history museums ...

, and the task of mounting the skeleton was given to

Wilhelm Gottlieb Tilesius Wilhelm Gottlieb Tilesius von Tilenau (17 July 1769 – 17 May 1857) was a German naturalist and explorer, physician, draftsman and engraver. He was a member of the Order of St. Vladimir and of the Legion of Honour. Early life and educatio ...

. This was one of the first attempts at reconstructing the skeleton of an extinct animal. Most of the reconstruction is correct, but Tilesius placed each tusk in the opposite socket, so that they curved outward instead of inward. The error was not corrected until 1899, and the correct placement of mammoth tusks was still a matter of debate into the 20th century. The 1901 excavation of the "Berezovka mammoth" is the best documented of the early finds. It was discovered at the Siberian Berezovka River (after a dog had noticed its smell), and the Russian authorities financed its excavation. The entire expedition took 10 months, and the specimen had to be cut to pieces before it could be transported to St. Petersburg. Most of the skin on the head as well as the trunk had been scavenged by predators, and most of the internal organs had rotted away. It was identified as a 35- to 40-year-old male, which had died 35,000 years ago. The animal still had grass between its teeth and on the tongue, showing that it had died suddenly. One of its shoulder blades was broken, which may have happened when it fell into a crevasse. It may have died of

asphyxiation Asphyxia or asphyxiation is a condition of deficient supply of oxygen to the body which arises from abnormal breathing. Asphyxia causes generalized hypoxia, which affects primarily the tissues and organs. There are many circumstances that ca ...

, as indicated by its erect penis. One third of a replica of the mammoth in the Museum of Zoology of

St. Petersburg Saint Petersburg ( rus, links=no, Санкт-Петербург, a=Ru-Sankt Peterburg Leningrad Petrograd Piter.ogg, r=Sankt-Peterburg, p=ˈsankt pʲɪtʲɪrˈburk), formerly known as Petrograd (1914–1924) and later Leningrad (1924–1991), i ...

is covered in skin and hair of the "Berezovka mammoth". By 1929, the remains of 34 mammoths with frozen soft tissues (skin, flesh, or organs) had been documented. Only four of them were relatively complete. Since then, about that many more have been found. In most cases, the flesh showed signs of decay before its freezing and later desiccation. Since 1860, Russian authorities have offered rewards of up to for finds of frozen woolly mammoth carcasses. Often, such finds were kept secret due to superstition. Several carcasses have been lost because they were not reported, and one was fed to dogs. Despite the rewards, native

Yakuts The Yakuts, or the Sakha ( sah, саха, ; , ), are a Turkic ethnic group who mainly live in the Republic of Sakha in the Russian Federation, with some extending to the Amur, Magadan, Sakhalin regions, and the Taymyr and Evenk Districts ...

were also reluctant to report mammoth finds to the authorities due to bad treatment of them in the past. In more recent years, scientific expeditions have been devoted to finding carcasses instead of relying solely on chance encounters. The most famous frozen specimen from Alaska is a calf nicknamed "Effie", which was found in 1948. It consists of the head, trunk, and a fore leg, and is about 25,000 years old.Lister, 2007. pp. 45–75 In 1977, the well-preserved carcass of a seven- to eight-month-old woolly mammoth calf named "Dima" was discovered. This carcass was recovered near a tributary of the

Kolyma River The Kolyma ( rus, Колыма, p=kəlɨˈma; sah, Халыма, translit=Khalyma) is a river in northeastern Siberia, whose basin covers parts of the Sakha Republic, Chukotka Autonomous Okrug, and Magadan Oblast of Russia. The Kolyma is fro ...

in northeastern Siberia. This specimen weighed about at death and was high and long. Radiocarbon dating determined that "Dima" died about 40,000 years ago. Its internal organs are similar to those of modern elephants, but its ears are only one-tenth the size of those of an African elephant of similar age. A less complete juvenile, nicknamed "Mascha", was found on the

Yamal Peninsula The Yamal Peninsula (russian: полуостров Ямал, poluostrov Yamal) is located in the Yamalo-Nenets Autonomous Okrug of northwest Siberia, Russia. It extends roughly 700 km (435 mi) and is bordered principally by the Kara ...

in 1988. It was 3–4 months old, and a laceration on its right foot may have been the cause of death. It is the westernmost frozen mammoth found. In 1997, a piece of mammoth tusk was discovered protruding from the tundra of the

in Siberia, Russia. In 1999, this 20,380-year-old carcass and 25 tons of surrounding sediment were transported by an

Mi-26 The Mil Mi-26 (russian: link=no, Миль Ми-26, NATO reporting name: Halo) is a Soviet/Russian heavy transport helicopter. Its product code is ''Izdeliye 90''. Operated by both military and civilian operators, it is the largest and most po ...

heavy lift helicopter to an ice cave in Khatanga. The specimen was nicknamed the " Jarkov mammoth". In October 2000, the careful defrosting operations in this cave began with the use of

hair dryer A hair dryer, hairdryer or blow dryer is an electromechanical device that blows ambient or hot air over damp hair to speed the evaporation of water to dry the hair. Blow dryers enable better control over the shape and style of hair, by accelerat ...

s to keep the hair and other soft tissues intact. In 2002, a well-preserved carcass was discovered near the Maxunuokha River in northern Yakutia, which was recovered during three excavations. This adult male specimen was called the "Yukagir mammoth", and is estimated to have lived around 18,560 years ago, and to have been 282.9 cm (9.2 ft) tall at the shoulder, and weighed between 4 and 5 tonnes. It is one of the best-preserved mammoths ever found due to the almost complete head, covered in skin, but without the trunk. Some postcranial remains were found, some with soft tissue. In 2007, the carcass of a female calf nicknamed "Lyuba" was discovered near the Yuribey River, where it had been buried for 41,800 years. By cutting a section through a molar and analysing its growth lines, they found that the animal had died at the age of one month. The mummified calf weighed , was high and in length. At the time of discovery, its eyes and trunk were intact and some fur remained on its body. Its organs and skin are very well preserved. "Lyuba" is believed to have been suffocated by mud in a river that its herd was crossing. After death, its body may have been colonised by bacteria that produce

lactic acid Lactic acid is an organic acid. It has a molecular formula . It is white in the solid state and it is miscible with water. When in the dissolved state, it forms a colorless solution. Production includes both artificial synthesis as well as nat ...

, which "pickled" it, preserving the mammoth in a nearly pristine state. In 2012, a juvenile was found in Siberia, which had man-made cut marks. Scientists estimated its age at death to be 2.5 years, and nicknamed it "

". Its skull and pelvis had been removed prior to discovery, but were found nearby. After being discovered, the skin of "Yuka" was prepared to produce a

taxidermy Taxidermy is the art of preserving an animal's body via mounting (over an armature) or stuffing, for the purpose of display or study. Animals are often, but not always, portrayed in a lifelike state. The word ''taxidermy'' describes the proc ...

mount. In 2019, a group of researchers managed to obtain signs of biological activity after transferring nuclei of "Yuka" into mouse

oocytes An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female g ...

. In 2013, a well-preserved carcass was found on Maly Lyakhovsky Island, one of the islands in the

New Siberian Islands The New Siberian Islands ( rus, Новосиби́рские Oстрова, r=Novosibirskiye Ostrova; sah, Саҥа Сибиир Aрыылара, translit=Saña Sibiir Arıılara) are an archipelago in the Extreme North of Russia, to the north o ...

archipelago, a female between 50 and 60 years old at the time of death. The carcass contained well-preserved muscular tissue. When it was extracted from the ice, liquid blood spilled from the abdominal cavity. The finders interpreted this as indicating woolly mammoth blood possessed antifreezing properties. In 2022, a complete female baby woolly mammoth was found by a miner in the Klondike gold fields of

, Canada. The specimen is estimated to have died 30.000 years ago, and was nicknamed "Nun cho ga", meaning "big baby animal" in the local

Hän language The Hän language (alternatively spelled as Haen) (also known as Dawson, Han-Kutchin, Moosehide; ISO 639-3 haa) is a Northern Athabaskan language spoken by the Hän Hwëch'in (translated to ''people who live along the river'', sometimes anglic ...

. It is the best preserved woolly mammoth

mummy A mummy is a dead human or an animal whose soft tissues and organs have been preserved by either intentional or accidental exposure to chemicals, extreme cold, very low humidity, or lack of air, so that the recovered body does not decay fu ...

found in North America, and was the same size as Lyuba.

Revival of the species

The existence of preserved soft tissue remains and DNA of woolly mammoths has led to the idea that the species could be resurrected by scientific means. Several methods have been proposed to achieve this.

Cloning Cloning is the process of producing individual organisms with identical or virtually identical DNA, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction. In the field of biotechnology, c ...

would involve removal of the DNA-containing

nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom * Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...

of the egg cell of a female elephant and replacement with a nucleus from woolly mammoth tissue. The cell would then be stimulated into dividing and inserted back into a female elephant. The resulting calf would have the genes of the woolly mammoth, although its fetal environment would be different. Most intact mammoths have had little usable DNA because of their conditions of preservation. There is not enough to guide the production of an embryo.Lister, 2007. pp. 42–43 A second method involves artificially inseminating an elephant egg cell with sperm cells from a frozen woolly mammoth carcass. The resulting offspring would be an elephant–mammoth hybrid, and the process would have to be repeated so more hybrids could be used in breeding. After several generations of cross-breeding these hybrids, an almost pure woolly mammoth would be produced. In one case, an Asian elephant and an African elephant produced a live calf named Motty, but it died of defects at less than two weeks old. The fact that sperm cells of modern mammals are viable for 15 years at most after deep-freezing makes this method unfeasible. Elephants by the water (7568684536)

Several projects are working on gradually replacing the genes in elephant cells with mammoth genes.The Long Now Foundation – Revive and Restore
. By 2015 and using the new CRISPR DNA editing technique, one team, led by George Church, had some woolly mammoth genes edited into the genome of an Asian elephant; focusing on cold-resistance initially, the target genes are for the external ear size, subcutaneous fat,

hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyt ...

, and hair attributes. If any method is ever successful, a suggestion has been made to introduce the hybrids to a wildlife reserve in Siberia called the

Pleistocene Park Pleistocene Park (russian: Плейстоценовый парк, Pleystotsenovyy park) is a nature reserve on the Kolyma River south of Chersky in the Sakha Republic, Russia, in northeastern Siberia, where an attempt is being made to re-crea ...

. Some researchers question the ethics of such recreation attempts. In addition to the technical problems, not much habitat is left that would be suitable for elephant-mammoth hybrids. Because the species was social and

gregarious Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...

, creating a few specimens would not be ideal. The time and resources required would be enormous, and the scientific benefits would be unclear, suggesting these resources should instead be used to preserve extant elephant species which are endangered. The ethics of using elephants as surrogate mothers in hybridisation attempts has been questioned, as most embryos would not survive, and knowing the exact needs of a hybrid elephant–mammoth calf would be impossible. Another concern is the introduction of unknown pathogens if de-extinction efforts were to succeed. In 2021, an Austin-based company raised funds to reintroduce the species in the

Arctic tundra In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless mou ...

Cultural significance

The woolly mammoth has remained culturally significant long after its extinction. Indigenous peoples of Siberia had long found what are now known to be woolly mammoth remains, collecting their tusks for the ivory trade. Native Siberians believed woolly mammoth remains to be those of giant mole-like animals that lived underground and died when burrowing to the surface. Woolly mammoth tusks had been articles of trade in Asia long before Europeans became acquainted with them. Güyük, the 13th-century Khan of the Mongols, is reputed to have sat on a throne made from mammoth ivory. Inspired by the Siberian natives' concept of the mammoth as an underground creature, it was recorded in the 16th-century Chinese pharmaceutical encyclopedia, '' Ben Cao Gangmu'', as ''yin shu'', "the hidden rodent". The

indigenous peoples of North America The Indigenous peoples of the Americas are the inhabitants of the Americas before the arrival of the European settlers in the 15th century, and the ethnic groups who now identify themselves with those peoples. Many Indigenous peoples of the Am ...

used woolly mammoth ivory and bone for tools and art. As in Siberia, North American natives had "myths of observation" explaining the remains of woolly mammoths and other elephants; the Bering Strait Inupiat believed the bones came from burrowing creatures, while other peoples associated them with primordial giants or "great beasts". Observers have interpreted legends from several Native American peoples as containing folk memory of extinct elephants, though other scholars are skeptical that folk memory could survive such a long time. Siberian mammoth ivory is reported to have been exported to Russia and Europe in the 10th century. The first Siberian ivory to reach western Europe was brought to London in 1611. When Russia occupied Siberia, the ivory trade grew and it became a widely exported commodity, with huge amounts being excavated. From the 19th century and onwards, woolly mammoth ivory became a highly prized commodity, used as raw material for many products. Today, it is still in great demand as a replacement for the now-banned export of elephant ivory, and has been referred to as "white gold". Local dealers estimate that 10 million mammoths are still frozen in Siberia, and conservationists have suggested that this could help save the living species of elephants from extinction. Elephants are hunted by poachers for their ivory, but if this could instead be supplied by the already extinct mammoths, the demand could instead be met by these. Trade in elephant ivory has been forbidden in most places following the 1989

Lausanne , neighboring_municipalities= Bottens, Bretigny-sur-Morrens, Chavannes-près-Renens, Cheseaux-sur-Lausanne, Crissier, Cugy, Écublens, Épalinges, Évian-les-Bains (FR-74), Froideville, Jouxtens-Mézery, Le Mont-sur-Lausanne, Lugrin (FR ...

Conference, but dealers have been known to label it as mammoth ivory to get it through customs. Mammoth ivory looks similar to elephant ivory, but the former is browner and the Schreger lines are coarser in texture. In the 21st century,

global warming In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...

has made access to Siberian tusks easier, since the permafrost thaws more quickly, exposing the mammoths embedded within it. Stories abound about frozen woolly mammoth meat that was consumed once defrosted, especially that of the "Berezovka mammoth", but most of these are considered dubious. The carcasses were in most cases decayed, and the stench so unbearable that only wild scavengers and the dogs accompanying the finders showed any interest in the flesh. Such meat apparently was once recommended against illness in China, and Siberian natives have occasionally cooked the meat of frozen carcasses they discovered. According to one of the more famous stories, members of

The Explorers Club The Explorers Club is an American-based international multidisciplinary professional society with the goal of promoting scientific exploration and field study. The club was founded in New York City in 1904, and has served as a meeting point fo ...

dined on meat of a frozen mammoth from Alaska in 1951. In 2016, a group of researchers genetically examined a sample of the meal, and found it to belong to a green sea turtle (it had also been claimed to belong to ''

Megatherium ''Megatherium'' ( ; from Greek () 'great' + () 'beast') is an extinct genus of ground sloths endemic to South America that lived from the Early Pliocene through the end of the Pleistocene. It is best known for the elephant-sized type species ' ...

''). The researchers concluded that the dinner had been a publicity stunt. In 2011, the Chinese palaeontologist Lida Xing

livestreamed Livestreaming is streaming media simultaneously recorded and broadcast in real-time over the internet. It is often referred to simply as streaming. Non-live media such as video-on-demand, vlogs, and YouTube videos are technically streamed, but no ...

while eating meat from a Siberian mammoth leg (thoroughly cooked and flavoured with salt) and told his audience it tasted bad and like soil. This triggered controversy and gained mixed reactions, but Xing stated he did it to promote science.

Alleged survival

There have been occasional claims that the woolly mammoth is not extinct and that small, isolated herds might survive in the vast and sparsely inhabited

tundra In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless mou ...

of the Northern Hemisphere. In the 19th century, several reports of "large shaggy beasts" were passed on to the Russian authorities by Siberian tribesmen, but no scientific proof ever surfaced. A French '' chargé d'affaires ''working in

Vladivostok Vladivostok ( rus, Владивосто́к, a=Владивосток.ogg, p=vɫədʲɪvɐˈstok) is the largest city and the administrative center of Primorsky Krai, Russia. The city is located around the Golden Horn Bay on the Sea of Japan, c ...

, M. Gallon, said in 1946 that in 1920, he had met a Russian fur-trapper who claimed to have seen living giant, furry "elephants" deep into the

taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruc ...

. Due to the large area of Siberia, the possibility that woolly mammoths survived into more recent times cannot be completely ruled out, but evidence indicates that they became extinct thousands of years ago. These natives likely had gained their knowledge of woolly mammoths from carcasses they encountered and that this is the source for their legends of the animal. In the late 19th century, rumours existed about surviving mammoths in Alaska. In 1899, Henry Tukeman detailed his killing of a mammoth in Alaska and his subsequent donation of the specimen to the

Smithsonian Institution The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums and education and research centers, the largest such complex in the world, created by the U.S. government "for the increase and diffusion of knowledge". Founded ...

Washington, DC ) , image_skyline = , image_caption = Clockwise from top left: the Washington Monument and Lincoln Memorial on the National Mall, United States Capitol, Logan Circle, Jefferson Memorial, White House, Adams Morgan ...

. The museum denied the story. The Swedish writer Bengt Sjögren suggested in 1962 that the myth began when the American biologist

Charles Haskins Townsend Charles Haskins Townsend (September 29, 1859 – January 28, 1944) was an American zoologist and naturalist who served as the director of the New York Aquarium, from 1902 to 1937. Early life The son of the Reverend Daniel W. Townsend and Elizabet ...

travelled in Alaska, saw

Inuit Inuit (; iu, ᐃᓄᐃᑦ 'the people', singular: Inuk, , dual: Inuuk, ) are a group of culturally similar indigenous peoples inhabiting the Arctic and subarctic regions of Greenland, Labrador, Quebec, Nunavut, the Northwest Territories ...

trading mammoth tusks, asked if mammoths were still living in Alaska, and provided them with a drawing of the animal.

Bernard Heuvelmans Bernard Heuvelmans (10 October 1916 – 22 August 2001) was a Belgian-French scientist, explorer, researcher, and writer probably best known, along with Scottish-American biologist Ivan T. Sanderson, as a founding figure in the pseudoscienc ...

included the possibility of residual populations of Siberian mammoths in his 1955 book, ''On The Track Of Unknown Animals''; while his book was a systematic investigation into possible unknown species, it became the basis of the

cryptozoology Cryptozoology is a pseudoscience and subculture that searches for and studies unknown, legendary, or extinct animals whose present existence is disputed or unsubstantiated, particularly those popular in folklore, such as Bigfoot, the Loch Ness ...

movement.

References

Bibliography

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External links

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Natural History Museum: "The last of the mammoths"
– three-minute video about the extinction of the woolly mammoth, presented by Adrian Lister
National Geographic: "Mammoth tusk treasure hunt"
– two-minute video about mammoth tusk collecting in modern Siberia {{DEFAULTSORT:Woolly Mammoth Mammoths Pleistocene proboscideans Pleistocene first appearances Pleistocene mammals of North America Holarctic fauna Holocene extinctions Extinct animals of Asia Extinct animals of the United States Extinct animals of Canada Extinct mammals of Europe Extinct mammals of North America Fossil taxa described in 1799 Cenozoic animals of North America Taxa named by Johann Friedrich Blumenbach Symbols of Alaska