The woolly mammoth (Mammuthus primigenius) is an extinct species of mammoth that lived from the Middle Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with the African Mammuthus subplanifrons in the early Pliocene. The woolly mammoth began to diverge from the steppe mammoth about 800,000 years ago in Siberia. Its closest extant relative is the Asian elephant. The Columbian mammoth (Mammuthus columbi) lived alongside the woolly mammoth in North America, and DNA studies show that the two hybridised with each other.

Woolly mammoth
Temporal range: Middle Pleistocene – Late Holocene 0.40–0.0037 Ma
Largest European specimen, a male at Südostbayerisches Naturkunde- und Mammut-Museum, Siegsdorf
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Genus: Mammuthus
M. primigenius
Binomial name
Mammuthus primigenius
(Blumenbach, 1799)
Dymaxion map showing the Late Pleistocene distribution of M. primigenius in blue (light blue was land at the time), inferred from fossil finds
  • Elephas primigenius Blumenbach, 1799
  • Elephas mammonteus Cuvier, 1799
  • Mammuthus boreus Brookes, 1828
  • Mammonteus primigenius Osborn, 1924
  • Elephas boreus Hay, 1924

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 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 2.67 and 3.49 m (8.8 and 11.5 ft) and weighed between 3.9 and 8.2 metric tons (4.3 and 9.0 short tons). Females reached 2.3–2.6 m (7.5–8.5 ft) in shoulder heights and weighed between 2.8–4 metric tons (3.1–4.4 short tons). A newborn calf weighed about 90 kg (200 lb). 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 and heat loss. It had long, curved tusks and four molars, 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. Individuals could probably reach the age of 60. Its habitat was 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 Late Pleistocene, with the last populations on mainland Siberia persisting until around 10,000 years ago, although isolated populations survived on St. Paul Island until 5,600 years ago, and on Wrangel Island until 4,000 years ago. After its extinction, humans continued using its ivory as a raw material, a tradition that continues today. With a genome project 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.


Copy of an interpretation of the "Adams mammoth" carcass from around 1800, with Johann Friedrich Blumenbach's handwriting

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 such as behemoths or giants. They were thought to be remains of modern elephants that had been brought to Europe during the Roman Republic, for example the war elephants of Hannibal and Pyrrhus of Epirus, or animals that had wandered north.[1] The first woolly mammoth remains studied by European scientists were examined by Hans Sloane in 1728 and consisted of fossilised teeth and tusks from Siberia. Sloane was the first to recognise that the remains belonged to elephants.[2] Sloane turned to another biblical explanation for the presence of elephants in the Arctic, asserting that they had been buried during the Great Flood, and that Siberia had previously been tropical before a drastic climate change.[3]

Others interpreted Sloane's conclusion slightly differently, arguing the flood had carried elephants from the tropics to the Arctic. 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.[4] In 1738, the German zoologist Johann Philipp Breyne 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.[5]

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.[1][6] Following Cuvier's identification, German naturalist Johann Friedrich Blumenbach gave the woolly mammoth its scientific name, Elephas primigenius, in 1799, placing it in the same genus as the Asian elephant (Elephas maximus). 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.[7] In 1828, the British naturalist Joshua Brookes used the name Mammuthus borealis for woolly mammoth fossils in his collection that he put up for sale, thereby coining a new genus name.[8]

1930s illustration of the lectotype molars; the left one is now lost.

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".[9] It may be a version of mehemot, the Arabic version of the biblical word "behemoth". Another possible origin is Estonian, where maa means "earth", and mutt means "mole". The word was first used in Europe during the early 17th century, when referring to maimanto tusks discovered in Siberia.[10] American president Thomas Jefferson, 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.[11]

By the early 20th century, the taxonomy of extinct elephants was complex. In 1942, American palaeontologist Henry Fairfield Osborn'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.[12] 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.[13]

Osborn chose two molars (found in Siberia and Osterode) from Blumenbach's collection at Göttingen University as the lectotype specimens for the woolly mammoth, since holotype 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 specimen in 1990. Resolutions to historical issues about the validity of the genus name Mammuthus and the type species designation of E. primigenius were also proposed.[14] 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.[7][15]


Georges Cuvier's 1796 comparison between the mandible of a woolly mammoth (bottom left and top right) and an Indian elephant (top left and bottom right)

The earliest known members of the Proboscidea, the clade which contains modern elephants, existed about 55 million years ago around the Tethys Sea. The closest known relatives of the Proboscidea are the sirenians (dugongs and manatees) and the hyraxes (an order of small, herbivorous mammals). The family Elephantidae existed 6 million years ago in Africa and includes the modern elephants and the mammoths. Among many now extinct clades, the mastodon (Mammut) is only a distant relative of the mammoths, and part of the separate family Mammutidae, which diverged 25 million years before the mammoths evolved.[16] The Asian elephant is the closest extant relative of the mammoths.[17] The following cladogram shows the placement of the woolly mammoth among Late Pleistocene and modern proboscideans, based on genetic data:[18][19]

Mammut americanum (American mastodon)

Notiomastodon platensis (South American gomphothere)


Mammuthus columbi (Columbian mammoth)

Mammuthus primigenius (woolly mammoth)

Elephas maximus (Asian elephant)

Palaeoloxodon antiquus (straight-tusked elephant)

Loxodonta cyclotis (African forest elephant)

Loxodonta africana (African bush elephant)

Comparison between the lower molars of a woolly mammoth (above) and a southern mammoth; note the lower number of enamel ridges in the older species (below)

Within six weeks from 2005–2006, three teams of researchers independently assembled mitochondrial genome profiles of the woolly mammoth from ancient DNA, which allowed them to confirm the close evolutionary relationship between mammoths and Asian elephants (Elephas maximus).[20][21] A 2015 DNA review confirmed Asian elephants as the closest living relative of the woolly mammoth.[22] 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.[23] 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.[24]

In 2008, much of the woolly mammoth's chromosomal DNA was mapped. The analysis showed that the woolly mammoth and the African elephant are 98.55% to 99.40% identical.[25] 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.[26] In 2012, proteins were confidently identified for the first time, collected from a 43,000-year-old woolly mammoth.[27]

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 molars; 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.[28][29] The short and tall skulls of woolly and Columbian mammoths (Mammuthus columbi) were the culmination of this process.[30]

The first known members of the genus Mammuthus are the African species Mammuthus subplanifrons from the Pliocene, and M. africanavus from the Pleistocene. 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.[28] Mammoths derived from M. trogontherii evolved molars with 26 ridges 400,000 years ago in Siberia and became the woolly mammoth. The earliest identified forms of woolly mammoth date to the Middle Pleistocene.[28] Woolly mammoths entered North America about 100,000 years ago by crossing the Bering Strait.[30]

Subspecies and hybridisation

Cast of an intermediate form between M. trogontherii and M. primigenius; M. p. fraasi, Staatliches Museum für Naturkunde Stuttgart
Specimen (formerly assigned to M. jeffersonii) suggested to be a hybrid between Columbian and woolly mammoths at the American Museum of Natural History

Individuals and populations showing transitional morphologies between each of the mammoth species are known, and primitive and derived species coexisted until the former disappeared. The different species and their intermediate forms have been termed "chronospecies". 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.[28] 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.[12][31][32]

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 hybrid between the two species.[33] 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.[34]

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. The study found that half of the ancestry of Columbian mammoths came from relatives of the Krestovka lineage (which probably represented the first mammoths that colonised the Americas) and the other half from the lineage of woolly mammoths, with the hybridisation happening more than 420,000 years ago, during the Middle Pleistocene. Later woolly and Columbian mammoths also interbred occasionally, and mammoth species may have hybridised routinely when brought together by glacial expansion. These findings were the first evidence of hybrid speciation 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 were not unique to woolly mammoths.[35][36]


Size of a large male (red) compared to a human and other mammoths

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. The average shoulder height for males of the species has been estimated at 2.8–3.15 metres (9.2–10.3 ft) with a weight of 4.5–6 tonnes (9,900–13,200 lb), with females being smaller like living elephants, with a shoulder height of 2.3–2.6 metres (7.5–8.5 ft) and a weight of 2.8–4 tonnes (6,200–8,800 lb).[37] This size is comparable to the largest living elephant species, the African bush elephant (Loxodonta africana), but is considerably smaller than the earlier Mammuthus meridionalis and Mammuthus trogontherii and the contemporary Mammuthus columbi.[37][38] The woolly mammoth exhibited size variation throughout its range, with individuals from Western Europe being considerably larger (with adult males estimated to be on average 2.99–3.31 metres (9.8–10.9 ft) tall and 5.2–6.9 tonnes (11,000–15,000 lb) in weight) than those found in Siberia (with adult males of this population being estimated on average 2.66–2.94 metres (8.7–9.6 ft) tall and 3.9–5.2 tonnes (8,600–11,500 lb) in weight). One of the largest recorded woolly mammoths is the Siegsdorf specimen from Germany, with an estimated shoulder height of 3.49 metres (11.5 ft) and an estimated body mass of 8.2 tonnes (18,000 lb).[38] A newborn calf would have weighed about 90 kilograms (200 lb).[39]

Size comparison of average-sized male and female woolly mammoths

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 girdle, since the opening that functions as the birth canal is always wider in females than in males.[40] 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".[41] The last woolly mammoth populations are claimed to have decreased in size and increased their sexual dimorphism, but this was dismissed in a 2012 study.[42]

Model at the Royal BC Museum

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 38 cm (15 in) long and 18–28 cm (7.1–11.0 in) across, and the ear of the 6- to 12-month-old frozen calf "Dima" was under 13 cm (5.1 in) long. The small ears reduced heat loss and frostbite, and the tail was short for the same reason, only 36 cm (14 in) 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 1.25 and 2.5 cm (0.49 and 0.98 in). They had a layer of fat up to 10 cm (3.9 in) 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.[43]

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 processes 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 10 cm (3.9 in) long, while the lower "thumb" was 5 cm (2.0 in) and was broader. The trunk of "Dima" was 76 cm (2.49 ft) long, whereas the trunk of the adult "Liakhov mammoth" was 2 metres (6.6 ft) long.[43] The well-preserved trunk of a juvenile specimen nicknamed "Yuka" 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.[44]


SEM magnified image of an overhair

The coat consisted of an outer layer of long, coarse "guard hair", which was 30 cm (12 in) on the upper part of the body, up to 90 cm (35 in) in length on the flanks and underside, and 0.5 mm (0.020 in) in diameter, and a denser inner layer of shorter, slightly curly under-wool, up to 8 cm (3.1 in) long and 0.05 mm (0.0020 in) in diameter. The hairs on the upper leg were up to 38 cm (15 in) long, and those of the feet were 15 cm (5.9 in) 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 60 cm (24 in) long, which were thicker than the guard hairs. The woolly mammoth likely moulted seasonally, and the heaviest fur was shed during spring.[45]

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.[45] This is thought to be for thermoregulation, helping them lose heat in their hot environments.[46] Comparison between the over-hairs of woolly mammoths and extant elephants show that they did not differ much in overall morphology.[47] Woolly mammoths had numerous sebaceous glands 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.[48]

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 pigmentation varied from hair to hair and within each hair.[43] A 2006 study sequenced the Mc1r gene (which influences hair colour in mammals) from woolly mammoth bones. Two alleles were found: a dominant (fully active) and a recessive (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.[49] A 2011 study showed that light individuals would have been rare.[50] 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.[51]


Skull from Poland with one broken and one downward spiralled tusk

Woolly mammoths had very long tusks (modified incisor teeth), which were more curved than those of modern elephants. The longest known male tusk is 4.05 m (13.3 ft) long (measured along the outside curve) and weighs 115.5 kg (255 lb), with a historical report of a 4.30 m (14.1 ft) long tusk found in Siberia, while the heaviest tusk is 121 kg (267 lb), suggested to have been 125–130 kg (276–287 lb) when complete.[52][53] 2.4–2.7 m (7.9–8.9 ft) and 45 kg (99 lb) was a more typical size. Female tusks were smaller and thinner, 1.5–1.8 m (4.9–5.9 ft) and weighing 9 kg (20 lb). For comparison, the record for longest tusks of the African bush elephant is 3.4 m (11 ft). 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 would occur than with straight tusks.[53][54][52]

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 2.5–15 cm (0.98–5.91 in) 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.[53][54]

Molar from Font de Champdamoy, France, Musée Georges-Garret

Woolly mammoths had four functional molar teeth at a time—two in the upper jaw and two in the lower. About 23 cm (9.1 in) of the crown was within the jaw, and 2.5 cm (1 in) 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 and dentine. 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, 1.3 cm (0.51 in), the third were 15 cm (6 in) 15 cm (5.9 in) long, and the sixth were about 30 cm (1 ft) long and weighed 1.8 kg (4 lb). The molars grew larger and contained more ridges with each replacement.[55] The woolly mammoth is considered to have had the most complex molars of any elephant.[54]


Life restoration of fauna during the Pleistocene epoch in northern Spain, by Mauricio Antón, 2004

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, cave hyenas, and large felines. The tusks may have been used in intraspecies fighting, such as fights over territory 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, 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.[56] 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.[43]

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.[57] 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 2 m (6.6 ft), and the juveniles ran to keep up.[58]

Woolly mammoth dental enamel from Poland has demonstrated that woolly mammoths were seasonally migratory. Recurring shifts in δ18O and 87Sr/86Sr found in layers of the enamel correspond to seasonal variations and indicate that Polish woolly mammoths inhabited southern Poland during winter but grazed the Polish midlands during summer.[59]

Adaptations to cold

Head and leg of the adult male "Yukagir mammoth" (the trunk is not preserved); note fur and small ears

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, 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.[44] As in reindeer and musk oxen, the haemoglobin 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.[60]

In a 2015 study, high-quality genome 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.[61][62]

A 2019 study of the woolly mammoth mitogenome suggest that these had metabolic adaptations related to extreme environments.[63] A genetic study from 2023 found that the woolly mammoth had already acquired a broad range of genes associated with the development of skin and hair, fat storage, metabolism, and the immune system by the time the species appeared, and that these continued to evolve within the last 700,000 years, including a gene that resulted in mammoths of the Late Quaternary having small ears.[64]


Mandibles and lower molars, Barcelona

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 plants, flowering plants, 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 tonnes would need to eat 180 kg (397 lb) 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 (Quaternary glaciation, 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 buds 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.[65] Isotope analysis shows that woolly mammoths fed mainly on C3 plants, unlike horses and rhinos.[66]

Scientists identified milk in the stomach and faecal matter in the intestines of the mammoth calf "Lyuba".[67] 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.[68] An isotope analysis of woolly mammoths from Yukon showed that the young nursed for at least 3 years, and were weaned 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.[69]

Male tusk with signs of wear

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 30 centimetres (12 in) 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.[70][71]

Life history

Cross sections of African elephant and woolly mammoth tusks; growth rings can be used to determine age

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 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.[72][73]

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 90 cm (35 in) 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.[72]

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 probably was from summer to autumn.[74] 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.[75]

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.[76] This feature indicates that, like bull elephants, male woolly mammoths entered "musth", a period of heightened aggressiveness. The glands are used especially by males to produce an oily substance with a strong smell called temporin. Their fur may have helped in spreading the scent further.[77] This was confirmed by a 2023 study that compared the testosterone level in the dentine of an adult African elephant tusk with that of a male woolly mammoth.[78]


Evidence of several different bone diseases has been found in woolly mammoths. The most common of these was osteoarthritis, found in 2% of specimens. One specimen from Switzerland had several fused vertebrae as a result of this condition. The "Yukagir mammoth" had suffered from spondylitis in two vertebrae, and osteomyelitis is known from some specimens. Several specimens have healed bone fractures, showing that the animals had survived these injuries.[79] Likewise, spondyloarthropathy has also been identified in woolly mammoth remains.[80] An extra number of cervical vertebrae has been found in 33% of specimens from the North Sea region, probably due to a drop in numbers and subsequent inbreeding.[81] Vertebral lesions in woolly mammoths have been speculated to have resulted from nutritional stress.[82] Parasitic flies and protozoa were identified in the gut of the calf "Dima".[83]

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, with half of these containing caries.[84] The teeth sometimes had cancerous growths.[85]

Distribution and habitat

The habitat of the woolly mammoth is known as "mammoth steppe" 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 of modern Russia, but the flora 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 areas at the time. The habitat of the woolly mammoth supported other grazing herbivores such as the woolly rhinoceros, wild horses, and bison.[84] The Altai-Sayan assemblages are the modern biomes most similar to the "mammoth steppe".[86] A 2014 study concluded that forbs (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.[87]

Mural depicting a herd walking near the Somme River in France, by Charles R. Knight, 1916

The southernmost woolly mammoth specimen known is from the Shandong province of China, and is 33,000 years old.[88] The southernmost European remains are from the Depression of Granada in Spain and are of roughly the same age.[89][90] DNA studies have helped determine the phylogeography 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.[91] 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), with Siberia being more uniformly cold and dry throughout the Late Pleistocene.[92] During the Younger Dryas age, woolly mammoths briefly expanded into north-east Europe, whereafter the mainland populations became extinct.[93]

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.[94] 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 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.[84]

Relationship with humans

Woolly mammoth engraved on ivory found in 1864, the first known contemporary depiction of a prehistoric animal

Modern humans co-existed with woolly mammoths during the Upper Palaeolithic 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 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 published his discovery of the Red Lady of Paviland 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 found an engraving of a woolly mammoth on a piece of mammoth ivory in the Abri de la Madeleine cave in Dordogne, 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.[95]

Various prehistoric depictions of woolly mammoths, including cave paintings (above) and sculptures

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 has the most depictions, 159, and some of the drawings are more than 2 metres (6.6 ft) in length. Other notable caves with mammoth depictions are the Chauvet Cave, Les Combarelles Cave, and Font-de-Gaume.[96] A depiction in the Cave of El Castillo may instead show Palaeoloxodon, the "straight-tusked elephant".[97]

"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 shaped as woolly mammoths have been found in France.[96] Some portable mammoth depictions may not have been produced where they were discovered, but could have moved around by ancient trading.[97]


Reconstructed bone hut based on finds in Mezhyrich, exhibited in Japan

Woolly mammoth bones were used as construction material for dwellings by both Neanderthals and modern humans during the ice age.[98] More than 70 such dwellings are known, mainly from the East European Plain. The bases of the huts were circular, and ranged from 8 to 24 square metres (86 to 258 sq ft). The arrangement of dwellings varied, and ranged from 1 to 20 m (3.3 to 65.6 ft) 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 40 cm (16 in) 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.[99]

Woolly mammoth ivory was used to create art objects. Several Venus figurines, including the Venus of Brassempouy and the Venus of Lespugue, were made from this material. Weapons made from ivory, such as daggers, spears, and a boomerang, 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.[100][71]

Artifacts made from woolly mammoth ivory; The Venus of Brassempouy, the Venus of Moravany, and the Lion-Man

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.[101]

At a site in southern Poland 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.[102] A specimen from the Mousterian age of Italy shows evidence of spear hunting by Neanderthals.[103] 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.[104] A site near the Yana River 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.[105] Two woolly mammoths from Wisconsin, the "Schaefer" and "Hebior mammoths", show evidence of having been butchered by Paleo-Indians.[106][107]


Most woolly mammoth populations disappeared during the late Pleistocene and mid-Holocene,[108] coinciding with the extinction of most North American Pleistocene megafauna (including the Columbian mammoth) as well as the extinctions or extirpations of steppe-associated fauna of Eurasia that coexisted with the mammoth species (such as the woolly rhinoceros, the cave lion, reindeer, saiga, the Arctic fox, and the steppe lemming). This extinction formed part of the Late Pleistocene extinctions, 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.[109][110]

Palaeolithic projectile points made from mammoth ivory, Pekárna cave

Whatever the cause, large mammals are generally more vulnerable than smaller ones due to their smaller population size and low reproduction rates.[111] Climatic patterns during the Last Interglacial (130–116 kyr BP) suggest that woolly mammoths and associated steppe faunas were sensitive to contractions of steppe-tundra habitats since they were adapted to cold, dry, and open environments. Genetic results and climatic models both indicate that habitats suitable for the woolly mammoth in Eurasia contracted during the interglacial period, which would have caused population bottleneck effects that restricted its range to a few northern areas. As the climate favoured colder environments, however, woolly mammoth populations rebounded during later glacial periods.[112]

The Last Glacial Period of the late Pleistocene is considered that of the maximum geographic distribution of the woolly mammoth, occupying most of Europe, northern Asia, and northern North America, although several barriers such as ice sheets, high mountain chains, deserts, year-round water surfaces, and other grasslands prevented them from spreading farther.[113] Towards the end of the Last Glacial period, from around 15,000 years ago, the mammoth steppe that the woolly mammoth inhabited was gradually replaced across most of Siberia with wet tundra and boreal and temperate forest, which the woolly mammoth would have found to be unfavourable habitat.[114]

Different woolly mammoth populations did not die out simultaneously across their range, but gradually became extinct over time.[115] The dynamics of different woolly mammoth populations varied as they experienced very different magnitudes of climatic and human impacts over time, suggesting that extinction causes would have varied by population.[116] Most populations disappeared between 14,000 and 10,000 years ago.[117] In Britain, woolly mammoths were still present between 14,500 and 14,000 BP.[118] The youngest fossils of the mainland population are from the Kyttyk Peninsula of Siberia and date to 9,650 years ago.[111][117]

Woolly mammoth and muskox remains displayed on Wrangel Island, where mammoths survived until 4,000 years ago

A small population of woolly mammoths survived on St. Paul Island, Alaska, well into the Holocene[119][120][121] with the most recently published date of extinction being 5,600 years B.P.[122] The last population known from fossils remained on Wrangel Island in the Arctic Ocean until 4,000 years ago, well into the start of human civilization and seveeral centures subsequent to the construction of the Great Pyramid of ancient Egypt.[123][124][125][126] 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 environmental DNA, mammoths survived in the Yukon until about 5,700 years ago, roughly concurrent with the St. Paul population, and on the Taymyr Peninsula 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.).[108][127] However, ancient environmental DNA in cold environments can be reworked from older sediments into younger sediments that clearly post-date extinction, raising doubt about validity of these dates.[128]

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.[129] 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[130] led to a 20%[129] to 30%[126] loss of heterozygosity, and a 65% loss in mitochondrial DNA diversity.[126] The population seems to have subsequently been stable, without suffering further significant loss of genetic diversity.[126][131] Genetic evidence thus implies the extinction of this final population was sudden, rather than the culmination of a gradual decline.[126]

Map showing climatic suitability for woolly mammoths in the Late Pleistocene and Holocene of Eurasia: red is increasing suitability, green is decreasing suitability. Black points are records of mammoths, black lines are the northern limit of humans

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.[132] It is not clear whether these genetic changes contributed to their extinction.[133] It has been proposed that these changes are consistent with the concept of genomic meltdown;[132] 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.[134]

The disappearance coincides roughly in time with the first evidence for humans on the island,[135] though other authors have suggested that woolly mammoths were almost certainly extinct for several centuries prior to the presence of humans on Wrangel Island (which dates to around 3,600 years ago).[114] The woolly mammoths of eastern Beringia (modern Alaska and Yukon) 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 proboscideans (mammoths, gomphotheres, and mastodons), as well as most of the rest of the megafauna, of the Americas.[136] 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,[136] perhaps in large measure as a result of a consequent reduction in the freshwater supply.[137]

Changes in climate shrank suitable mammoth habitat from 7,700,000 km2 (3,000,000 sq mi) 42,000 years ago to 800,000 km2 (310,000 sq mi) by 6,000 years ago.[138][139] Woolly mammoths survived an even greater loss of habitat at the end of the Penultimate Glacial Period and onset of the Last Interglacial approximately 125,000 years ago.[140][141] Studies of an 11,300–11,000-year-old trackway 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.[58] It has been suggested that human hunting exerted a significant pressure on woolly mammoth populations for thousands of years across their range, making the population abundance of woolly mammoths considerably lower than it would have been otherwise even prior to their range decline, and likely hastened the range collapse of woolly mammoths in response to climate change.[116]

The decline of the woolly mammoth could have increased temperatures by up to 0.2 °C (0.36 °F) 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.[142]

Fossil specimens

Mounted "family group" from Tomsk

Woolly mammoth fossils have been found in many different types of deposits, including former rivers and lakes, and in "Doggerland" in the North Sea, 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' graveyards", 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.[143]

Skull discovered by fishermen in the North Sea ("Doggerland"), at Celtic and Prehistoric Museum, Ireland

Apart from frozen remains, the only soft tissue known is from a specimen that was preserved in a petroleum seep in Starunia, Poland. Frozen remains of woolly mammoths have been found in the northern parts of Siberia 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.[144]

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.[145] 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 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.[146]

Frozen specimens

The "Adams mammoth" as illustrated in the 1800s (left) and on exhibit in Vienna; skin can be seen on its head and feet.

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.[147] 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.[148] While in Yakutsk in 1806, Michael Friedrich Adams 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, and the task of mounting the skeleton was given to Wilhelm Gottlieb Tilesius.[4][149] 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.[150][151]

The "Berezovka mammoth" during excavation in 1901 (left), and a model partially covered by its skin, Museum of Zoology in St. Petersburg

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, as indicated by its erect penis. One third of a replica of the mammoth in the Museum of Zoology of St. Petersburg is covered in skin and hair of the "Berezovka mammoth".[144][145]

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.[152] Since 1860, Russian authorities have offered rewards of up to 1000 ₽ 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.[143] Despite the rewards, native Yakuts were also reluctant to report mammoth finds to the authorities due to bad treatment of them in the past.[147] 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.[143]

"Dima", a frozen calf, during excavation (left), and as exhibited in the Museum of Zoology; note fur on the legs

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 in northeastern Siberia. This specimen weighed about 100 kg (220 lb) at death and was 104 cm (41 in) high and 115 cm (45 in) 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 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.[153]

In 1997, a piece of mammoth tusk was discovered protruding from the tundra of the Taymyr Peninsula in Siberia, Russia. In 1999, this 20,380-year-old carcass and 25 tons of surrounding sediment were transported by an Mi-26 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 dryers to keep the hair and other soft tissues intact.[154][155]

The calf "Lyuba", in Royal BC Museum and IFC Mall

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.[76]

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.[68][156] 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.[75] The mummified calf weighed 50 kg (110 lb), was 85 cm (33 in) high and 130 cm (51 in) in length.[157][158] 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.[159] "Lyuba" is believed to have been suffocated by mud in a river that its herd was crossing.[68][160] After death, its body may have been colonised by bacteria that produce lactic acid, which "pickled" it, preserving the mammoth in a nearly pristine state.[68]

The frozen calf "Yuka" (left), and its skull and jaw which may have been extracted from the carcass by prehistoric humans

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 "Yuka". Its skull and pelvis had been removed prior to discovery, but were found nearby.[104][161] After being discovered, the skin of "Yuka" was prepared to produce a taxidermy mount.[44] In 2019, a group of researchers managed to obtain signs of biological activity after transferring nuclei of "Yuka" into mouse oocytes.[162]

In 2013, a well-preserved carcass was found on Maly Lyakhovsky Island, one of the islands in the New Siberian Islands 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.[163] In 2022, a complete female baby woolly mammoth was found by a miner in the Klondike gold fields of Yukon, 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. It is the best preserved woolly mammoth mummy found in North America, and was the same size as Lyuba.[164][165]

Possible revival

Models of an adult and the calf "Dima" in State Museum of Natural History Stuttgart

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 would involve removal of the DNA-containing nucleus 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.[166][167]

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. The fact that sperm cells of modern mammals are viable for 15 years at most after deep-freezing makes this method unfeasible.[167]

Elephants are highly gregarious, as shown by these Sri Lankan elephants

Several projects are working on gradually replacing the genes in elephant cells with mammoth genes.[168][169] 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,[170] the target genes are for the external ear size, subcutaneous fat, hemoglobin, and hair attributes.[171][172] 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.[173]

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, 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.[167][174][175] 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.[176] Another concern is the introduction of unknown pathogens if de-extinction efforts were to succeed.[177] In 2021, an Austin-based company raised funds to reintroduce the species in the Arctic tundra.[178]

Cultural significance

A mammoth tusk with Inuit carvings of scenes on the Yukon River, 19th century, De Young Museum

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.[179][180] 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.[147] 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".[181]Patkanov 1897, pp. 123–124

The indigenous peoples of North America used woolly mammoth ivory and bone for tools and art.[182] 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".[183][184][185] 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.[183][185][186]

Peter III and Elizabeth of Russia carved in mammoth ivory

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".[187]

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 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.[187] In the 21st century, global warming has made access to Siberian tusks easier, since the permafrost thaws more quickly, exposing the mammoths embedded within it.[188]

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.[189] According to one of the more famous stories, members of The Explorers Club 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). The researchers concluded that the dinner had been a publicity stunt.[190] In 2011, the Chinese palaeontologist Lida Xing livestreamed 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.[191] In 2023, An Australian cultured meat start-up, Vow, revealed a lab-grown "mammoth meatball" produced using a DNA sequence from the woolly mammoth. The meatball sparked conversations about the potential of cultured meat as a sustainable food source, highlighting its environmental benefits compared to traditional agriculture.[192]

Alleged survival

Woolly mammoths represented on the coats of arms of regions in Russia and Ukraine

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 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, 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.[193] 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.[194]

In the late 19th century, rumours existed about surviving mammoths in Alaska.[193] In 1899, Henry Tukeman detailed his killing of a mammoth in Alaska and his subsequent donation of the specimen to the Smithsonian Institution in Washington, DC. The museum denied the story.[195] The Swedish writer Bengt Sjögren suggested in 1962 that the myth began when the American biologist Charles Haskins Townsend travelled in Alaska, saw Inuit trading mammoth tusks, asked if mammoths were still living in Alaska, and provided them with a drawing of the animal.[193] Bernard Heuvelmans 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 movement.[196]


  1. ^ a b Switek, B. (2010). Written in Stone: Evolution, the Fossil Record, and Our Place in Nature. Bellevue Literary Press. pp. 174–180. ISBN 978-1-934137-29-1.
  2. ^ Sloane, H. (1727–1728). "An Account of Elephants Teeth and Bones Found under Ground". Philosophical Transactions. 35 (399–406): 457–471. Bibcode:1727RSPT...35..457S. doi:10.1098/rstl.1727.0042.
  3. ^ Sloane, H. (1727–1728). "Of Fossile Teeth and Bones of Elephants. Part the Second". Philosophical Transactions. 35 (399–406): 497–514. Bibcode:1727RSPT...35..497S. doi:10.1098/rstl.1727.0048.
  4. ^ a b The Academy of Natural Sciences (2007). "Woolly Mammoth (Mammuthus primigenius)". The Academy of Natural Sciences. Archived from the original on 27 September 2007. Retrieved 29 September 2007.
  5. ^ Breyne, J. P.; s., T.; Wolochowicz, M. (1737). "A Letter from John Phil. Breyne, M. D. F. R. S. To Sir Hans Sloane, Bart. Pres. R. S. With Observations, and a Description of Some Mammoth's Bones Dug up in Siberia, Proving Them to Have Belonged to Elephants". Philosophical Transactions of the Royal Society of London. 40 (445–451): 124–138. Bibcode:1737RSPT...40..124P. doi:10.1098/rstl.1737.0026.
  6. ^ Cuvier, G. (1796). "Mémoire sur les épèces d'elephans tant vivantes que fossils, lu à la séance publique de l'Institut National le 15 germinal, an IV". Magasin Encyclopédique, 2e Anée (in French): 440–445.
  7. ^ a b Reich, M.; Gehler, A.; Mohl, D.; van der Plicht, H.; Lister, A. M. (2007). "The rediscovery of type material of Mammuthus primigenius (Mammalia: Proboscidea)". International Mammoth Conference IV (Poster): 295.
  8. ^ Brookes, J. (1828). A catalogue of the anatomical & zoological museum of Joshua Brookes. Vol. 1. London: Richard Taylor. p. 73. Archived from the original on 24 September 2015.
  9. ^ "Mammoth entry in Oxford English Dictionary". 2000.
  10. ^ Lister, 2007. p. 49
  11. ^ Simpson, J. (2009). "Word Stories: Mammoth Archived 22 May 2013 at the Wayback Machine." Oxford English Dictionary Online, Oxford University Press. Accessed 5 June 2009.
  12. ^ a b Osborn, H. F. (1942). Percy, M. R. (ed.). Proboscidea: A monograph of the discovery, evolution, migration and extinction of the mastodonts and elephants of the world. Vol. 2. New York: J. Pierpont Morgan Fund. pp. 1116–1169. Archived from the original on 13 March 2016.
  13. ^ Maglio, V. J. (1973). "Origin and evolution of the Elephantidae". Transactions of the American Philosophical Society. 63 (3): 1–149. doi:10.2307/1379357. JSTOR 1379357.
  14. ^ Garutt, W. E.; Gentry, A.; Lister, A. M. (1990). "Mammuthus Brookes, 1828 (Mammalia, Proboscidea): proposed conservation, and Elephas primigenius Blumenbach, 1799 (currently Mammuthus primigenius): proposed designation as the type species of Mammuthus, and designation of a neotype". Bulletin of Zoological Nomenclature. 47: 38–44. doi:10.5962/bhl.part.2651. Archived from the original on 13 July 2015.
  15. ^ Reich, M.; Gehler, A. (2008). "Giants' Bones and Unicorn Horns Ice Age Elephants Offer 21st Century Insights". Collections – Wisdom, Insight, Innovation. 8: 44–50.
  16. ^ Lister, 2007. pp. 18–21
  17. ^ Shoshani, J.; Ferretti, M. P.; Lister, A. M.; Agenbroad, L. D.; Saegusa, H.; Mol, D.; Takahashi, K. (2007). "Relationships within the Elephantinae using hyoid characters". Quaternary International. 169–170: 174–185. Bibcode:2007QuInt.169..174S. doi:10.1016/j.quaint.2007.02.003.
  18. ^ Palkopoulou, Eleftheria; Lipson, Mark; Mallick, Swapan; Nielsen, Svend; Rohland, Nadin; Baleka, Sina; Karpinski, Emil; Ivancevic, Atma M.; To, Thu-Hien; Kortschak, R. Daniel; Raison, Joy M. (2018). "A comprehensive genomic history of extinct and living elephants". Proceedings of the National Academy of Sciences. 115 (11): E2566–E2574. Bibcode:2018PNAS..115E2566P. doi:10.1073/pnas.1720554115. ISSN 0027-8424. PMC 5856550. PMID 29483247.
  19. ^ Baleka, Sina; Varela, Luciano; Tambusso, P. Sebastián; Paijmans, Johanna L.A.; Mothé, Dimila; Stafford, Thomas W.; Fariña, Richard A.; Hofreiter, Michael (January 2022). "Revisiting proboscidean phylogeny and evolution through total evidence and palaeogenetic analyses including Notiomastodon ancient DNA". iScience. 25 (1): 103559. doi:10.1016/j.isci.2021.103559. PMC 8693454. PMID 34988402.
  20. ^ Gross, L. (2006). "Reading the Evolutionary History of the Woolly Mammoth in Its Mitochondrial Genome". PLOS Biology. 4 (3): e74. doi:10.1371/journal.pbio.0040074. PMC 1360100. PMID 20076539.
  21. ^ Cooper, A. (2006). "The year of the mammoth". PLOS Biology. 4 (3): e78. doi:10.1371/journal.pbio.0040078. PMC 1360097. PMID 16448215.
  22. ^ Roca, Alfred L.; Ishida, Yasuko; Brandt, Adam L.; Benjamin, Neal R.; Zhao, Kai; Georgiadis, Nicholas J. (2015). "Elephant Natural History: A Genomic Perspective". Annual Review of Animal Biosciences. 3 (1): 139–167. doi:10.1146/annurev-animal-022114-110838. PMID 25493538.
  23. ^ Krause, J.; Dear, P. H.; Pollack, J. L.; Slatkin, M.; Spriggs, H.; Barnes, I.; Lister, A. M.; Ebersberger, I.; Pääbo, S.; Hofreiter, M. (2005). "Multiplex amplification of the mammoth mitochondrial genome and the evolution of Elephantidae". Nature. 439 (7077): 724–727. Bibcode:2006Natur.439..724K. doi:10.1038/nature04432. PMID 16362058. S2CID 4318327.
  24. ^ Rohland, N.; Reich, D.; Mallick, S.; Meyer, M.; Green, R. E.; Georgiadis, N. J.; Roca, A. L.; Hofreiter, M. (2010). Penny, David (ed.). "Genomic DNA Sequences from Mastodon and Woolly Mammoth Reveal Deep Speciation of Forest and Savanna Elephants". PLOS Biology. 8 (12): e1000564. doi:10.1371/journal.pbio.1000564. PMC 3006346. PMID 21203580.
  25. ^ Will findings recreate the woolly mammoth? Archived 11 February 2009 at the Wayback Machine, Pittsburgh Post-Gazette, 20 November 2008
  26. ^ "Woolly-Mammoth Genome Sequenced". Science Daily. 20 November 2008. Archived from the original on 11 January 2011. Retrieved 22 June 2010.
  27. ^ Cappellini, E.; Jensen, L. J.; Szklarczyk, D.; Ginolhac, A. L.; Da Fonseca, R. A. R.; Stafford, T. W.; Holen, S. R.; Collins, M. J.; Orlando, L.; Willerslev, E.; Gilbert, M. T. P.; Olsen, J. V. (2012). "Proteomic analysis of a Pleistocene mammoth femur reveals more than one hundred ancient bone proteins". Journal of Proteome Research. 11 (2): 917–926. doi:10.1021/pr200721u. PMID 22103443.
  28. ^ a b c d Lister, A. M.; Sher, A. V.; Van Essen, H.; Wei, G. (2005). "The pattern and process of mammoth evolution in Eurasia" (PDF). Quaternary International. 126–128: 49–64. Bibcode:2005QuInt.126...49L. doi:10.1016/j.quaint.2004.04.014.
  29. ^ Ferretti, M. P. (2003). "Structure and evolution of mammoth molar enamel". Acta Palaeontologica Polonica. 3. 48: 383–396.
  30. ^ a b Lister, 2007. pp. 12–43
  31. ^ Foronova, I. V.; Zudin, A. N. (2001). Discreteness of evolution and variability in mammoth lineage: method for group study (PDF). The World of Elephants – Proceedings of the 1st International Congress. Rome. pp. 540–543. Archived (PDF) from the original on 24 October 2014.
  32. ^ Foronova, I.V. (2014). "Mammuthus intermedius (Proboscidea, Elephantidae) from the late Middle Pleistocene of the southern Western and Central Siberia, Russia: the problem of intermediate elements in the mammoth lineage" (PDF). Russian Journal of Theriology. 2. 13 (2): 71–82. doi:10.15298/rusjtheriol.13.2.03. Archived (PDF) from the original on 4 March 2016.
  33. ^ Enk, J.; Devault, A.; Debruyne, R.; King, C. E.; Treangen, T.; O'Rourke, D.; Salzberg, S. L.; Fisher, D.; MacPhee, R.; Poinar, H. (2011). "Complete Columbian mammoth mitogenome suggests interbreeding with woolly mammoths". Genome Biology. 12 (5): R51. doi:10.1186/gb-2011-12-5-r51. PMC 3219973. PMID 21627792.
  34. ^ Lister, A. M.; Sher, A. V. (13 November 2015). "Evolution and dispersal of mammoths across the Northern Hemisphere". Science. 350 (6262): 805–809. Bibcode:2015Sci...350..805L. doi:10.1126/science.aac5660. PMID 26564853. S2CID 206639522.
  35. ^ van der Valk, T.; Pečnerová, P.; Díez-del-Molino, D.; Bergström, A.; Oppenheimer, J.; Hartmann, S.; Xenikoudakis, G.; Thomas, J. A.; Dehasque, M.; Sağlıcan, E.; Fidan, F. Rabia; Barnes, I.; Liu, S.; Somel, M.; Heintzman, P. D.; Nikolskiy, P.; Shapiro, B.; Skoglund, P.; Hofreiter, M.; Lister, A. M.; Götherström, A.; Dalén, L. (2021). "Million-year-old DNA sheds light on the genomic history of mammoths". Nature. 591 (7849): 265–269. Bibcode:2021Natur.591..265V. doi:10.1038/s41586-021-03224-9. ISSN 1476-4687. PMC 7116897. PMID 33597750.
  36. ^ Callaway, E. (2021). "Million-year-old mammoth genomes shatter record for oldest ancient DNA". Vol. 590, no. 7847. pp. 537–538. doi:10.1038/d41586-021-00436-x. Retrieved 17 February 2021.
  37. ^ a b Larramendi, Asier; Palombo, Maria Rita; Marano, Federica (2017). "Reconstructing the life appearance of a Pleistocene giant: size, shape, sexual dimorphism and ontogeny of Palaeoloxodon antiquus (Proboscidea: Elephantidae) from Neumark-Nord 1 (Germany)" (PDF). Bollettino della Società Paleontologica Italiana (3): 299–317. doi:10.4435/BSPI.2017.29. ISSN 0375-7633. Archived from the original (PDF) on 30 September 2023.
  38. ^ a b Larramendi, Asier (2015). "Proboscideans: Shoulder Height, Body Mass and Shape". Acta Palaeontologica Polonica. 61. doi:10.4202/app.00136.2014.
  39. ^ Lister, 2007. pp. 82–87
  40. ^ Lister, 2007. pp. 174–175
  41. ^ Vartanyan, S. L.; Arslanov, K. A.; Karhu, J. A.; Possnert, G. R.; Sulerzhitsky, L. D. (2008). "Collection of radiocarbon dates on the mammoths (Mammuthus primigenius) and other genera of Wrangel Island, northeast Siberia, Russia" (PDF). Quaternary Research. 70 (1): 51–59. Bibcode:2008QuRes..70...51V. doi:10.1016/j.yqres.2008.03.005. S2CID 111383180.
  42. ^ Den Ouden, N.; Reumer, J. W. F.; Van Den Hoek Ostende, L. W. (2012). "Did mammoth end up a lilliput? Temporal body size trends in Late Pleistocene Mammoths, Mammuthus primigenius (Blumenbach, 1799) inferred from dental data". Quaternary International. 255: 53–58. Bibcode:2012QuInt.255...53D. doi:10.1016/j.quaint.2011.07.038.
  43. ^ a b c d Lister, 2007. pp. 82–87
  44. ^ a b c Plotnikov, V. V.; Maschenko, E. N.; Pavlov, I. S.; Protopopov, A. V.; Boeskorov, G. G.; Petrova, E. A. (2015). "New data on trunk morphology in the woolly mammoth, Mammuthus primigenius (Blumenbach)". Paleontological Journal. 49 (2): 200–210. Bibcode:2015PalJ...49..200P. doi:10.1134/S0031030115020070. S2CID 84849714.
  45. ^ a b Lister, 2007. pp. 83–84
  46. ^ Myhrvold, C. L.; Stone, H. A.; Bou-Zeid, E. (10 October 2012). "What is the use of elephant hair?". PLOS ONE. 7 (10): e47018. Bibcode:2012PLoSO...747018M. doi:10.1371/journal.pone.0047018. PMC 3468452. PMID 23071700.
  47. ^ Valente, A. (1983). "Hair structure of the woolly mammoth, Mammuthus primigenius and the modern elephants, Elephas maximus and Loxodonta africana". Journal of Zoology. 199 (2): 271–274. doi:10.1111/j.1469-7998.1983.tb02095.x.
  48. ^ Repin, V. E.; Taranov, O. S.; Ryabchikova, E. I.; Tikhonov, A. N.; Pugachev, V. G. (2004). "Sebaceous Glands of the Woolly Mammoth, Mammothus primigenius Blum.: Histological Evidence". Doklady Biological Sciences. 398 (1–6): 382–384. doi:10.1023/B:DOBS.0000046662.43270.66. PMID 15587793. S2CID 6401669.
  49. ^ Rompler, H.; Rohland, N.; Lalueza-Fox, C.; Willerslev, E.; Kuznetsova, T.; Rabeder, G.; Bertranpetit, J.; Schöneberg, T.; Hofreiter, M. (2006). "Nuclear Gene Indicates Coat-Color Polymorphism in Mammoths" (PDF). Science. 313 (5783): 62. doi:10.1126/science.1128994. PMID 16825562. S2CID 20153467.
  50. ^ Workman, C.; Dalen, L.; Vartanyan, S.; Shapiro, B.; Kosintsev, P.; Sher, A.; Gotherstrom, A.; Barnes, I. (2011). "Population-level genotyping of coat colour polymorphism in woolly mammoth (Mammuthus primigenius)". Quaternary Science Reviews. 30 (17–18): 2304–2308. Bibcode:2011QSRv...30.2304W. doi:10.1016/j.quascirev.2010.08.020.
  51. ^ Tridico, Silvana R.; Rigby, Paul; Kirkbride, K. Paul; Haile, James; Bunce, Michael (2014). "Megafaunal split ends: microscopical characterisation of hair structure and function in extinct woolly mammoth and woolly rhino". Quaternary Science Reviews. 83: 68–75. Bibcode:2014QSRv...83...68T. doi:10.1016/j.quascirev.2013.10.032. Archived from the original on 2 November 2017.
  52. ^ a b Boeskorov, G.; Tikhonov, A.; Shchelchkova, M.; Ballard, J. P.; Mol, D. (2020). "Big tuskers: Maximum sizes of tusks in woolly mammoths - Mammuthus primigenius (Blumenbach) - from East Siberia". Quaternary International. 537: 88–96. Bibcode:2020QuInt.537...88B. doi:10.1016/j.quaint.2019.12.023. S2CID 213262363.
  53. ^ a b c Lister, 2007. pp. 94–95
  54. ^ a b c Kurten, B.; Anderson, E. (1980). Pleistocene Mammals of North America. New York: Columbia University Press. pp. 348–354. ISBN 978-0-231-03733-4.
  55. ^ Lister, 2007. pp. 92–93
  56. ^ Lister, 2007. pp. 95–105
  57. ^ Lister, 2007. pp. 62–63
  58. ^ a b McNeil, P.; Hills, L.; Kooyman, B.; Tolman, S. (2005). "Mammoth tracks indicate a declining Late Pleistocene population in southwestern Alberta, Canada". Quaternary Science Reviews. 24 (10–11): 1253–1259. Bibcode:2005QSRv...24.1253M. doi:10.1016/j.quascirev.2004.08.019.
  59. ^ Kowalik, Nina; Anczkiewicz, Robert; Müller, Wolfgang; Spötl, Christoph; Bondioli, Luca; Nava, Alessia; Wojtal, Piotr; Wilczyński, Jarosław; Koziarska, Marta; Matyszczak, Milena (15 April 2023). "Revealing seasonal woolly mammoth migration with spatially-resolved trace element, Sr and O isotopic records of molar enamel". Quaternary Science Reviews. 306: 108036. doi:10.1016/j.quascirev.2023.108036. Retrieved 3 May 2024 – via Elsevier Science Direct.
  60. ^ Campbell, K. L.; Roberts, J. E. E.; Watson, L. N.; Stetefeld, J. R.; Sloan, A. M.; Signore, A. V.; Howatt, J. W.; Tame, J. R. H.; Rohland, N.; Shen, T. J.; Austin, J. J.; Hofreiter, M.; Ho, C.; Weber, R. E.; Cooper, A. (2010). "Substitutions in woolly mammoth hemoglobin confer biochemical properties adaptive for cold tolerance". Nature Genetics. 42 (6): 536–540. doi:10.1038/ng.574. PMID 20436470. S2CID 9670466.
  61. ^ Lynch, V.; Bedoya-Reina, O. C.; Ratan, A.; Sulak, M.; Drautz-Moses, D. I.; Perry, G. H.; Miller, W.; Schuster, S. C. (2015). "Elephantid genomes reveal the molecular bases of Woolly Mammoth adaptations to the arctic". Cell Reports. 12 (2): 217–228. doi:10.1016/j.celrep.2015.06.027. hdl:10220/38768. PMID 26146078.
  62. ^ Ewen Callaway, Nature magazine (4 May 2015). "Mammoth Genomes Provide Recipe for Creating Arctic Elephants". Archived from the original on 5 May 2015.
  63. ^ Ngatia, J. N.; Lan, T. M.; Dinh, T. D.; Zhang, L.; Ahmed, Ahmed Khalid; Xu, Yan Chun (2019). "Signals of positive selection in mitochondrial protein-coding genes of woolly mammoth: Adaptation to extreme environments?". Ecology and Evolution. 9 (12): 6821–6832. doi:10.1002/ece3.5250. PMC 6662336. PMID 31380018.
  64. ^ Díez-del-Molino, David; Dehasque, Marianne; Chacón-Duque, J. Camilo; Pečnerová, Patrícia; Tikhonov, Alexei; Protopopov, Albert; Plotnikov, Valeri; Kanellidou, Foteini; Nikolskiy, Pavel; Mortensen, Peter; Danilov, Gleb K.; Vartanyan, Sergey; Gilbert, M. Thomas P.; Lister, Adrian M.; Heintzman, Peter D.; van der Valk, Tom; Dalén, Love (2023). "Genomics of adaptive evolution in the woolly mammoth". Current Biology. 33 (9): 1753–1764.e4. doi:10.1016/j.cub.2023.03.084. PMID 37030294.
  65. ^ Lister, 2007. pp. 88–91
  66. ^ Bocherens, H.; Fizet, M.; Mariotti, A.; Gangloff, R. A.; Burns, J. A. (1994). "Contribution of isotopic biogeochemistry (13C,15N,18O) to the paleoecology of mammoths (Mammuthus primigenius)". Historical Biology. 7 (3): 187–202. Bibcode:1994HBio....7..187B. doi:10.1080/10292389409380453.
  67. ^ Van Geel, B.; Fisher, D. C.; Rountrey, A. N.; Van Arkel, J.; Duivenvoorden, J. F.; Nieman, A. M.; Van Reenen, G. B. A.; Tikhonov, A. N.; Buigues, B.; Gravendeel, B. (2011). "Palaeo-environmental and dietary analysis of intestinal contents of a mammoth calf (Yamal Peninsula, northwest Siberia)". Quaternary Science Reviews. 30 (27–28): 3935–3946. Bibcode:2011QSRv...30.3935V. doi:10.1016/j.quascirev.2011.10.009.
  68. ^ a b c d Fisher, D. C.; Tikhonov, A. N.; Kosintsev, P. A.; Rountrey, A. N.; Buigues, B.; Van Der Plicht, J. (2012). "Anatomy, death, and preservation of a woolly mammoth (Mammuthus primigenius) calf, Yamal Peninsula, northwest Siberia" (PDF). Quaternary International. 255: 94–105. Bibcode:2012QuInt.255...94F. doi:10.1016/j.quaint.2011.05.040. hdl:11370/a3961dcc-4eaf-47fb-9ad7-904d79a0f4f8. S2CID 35667021.
  69. ^ Metcalfe, J. Z.; Longstaffe, F. J.; Zazula, G. D. (2010). "Nursing, weaning, and tooth development in woolly mammoths from Old Crow, Yukon, Canada: Implications for Pleistocene extinctions". Palaeogeography, Palaeoclimatology, Palaeoecology. 298 (3–4): 257–270. Bibcode:2010PPP...298..257M. doi:10.1016/j.palaeo.2010.09.032.
  70. ^ Lister, 2007. pp. 92–95
  71. ^ a b Pfeifer, S. J.; Hartramph, W. L.; Kahlke, R.-D.; Müller, F. A. (2019). "Mammoth ivory was the most suitable osseous raw material for the production of Late Pleistocene big game projectile points". Scientific Reports. 9 (1): 2303. doi:10.1038/s41598-019-38779-1. PMC 6381109. PMID 30783179.
  72. ^ a b Lister, 2007. pp. 83–107.
  73. ^ Herbert, B.; Fisher, D. (5 May 2010). "A Mammoth Find: Clues to the Past, Present and Future". Helix. Northwestern University. Archived from the original on 18 August 2016. Retrieved 1 August 2016.
  74. ^ Lister, 2007. pp. 104–105
  75. ^ a b Rountrey, A. N.; Fisher, D. C.; Tikhonov, A. N.; Kosintsev, P. A.; Lazarev, P. A.; Boeskorov, G.; Buigues, B. (2012). "Early tooth development, gestation, and season of birth in mammoths". Quaternary International. 255: 196–205. Bibcode:2012QuInt.255..196R. doi:10.1016/j.quaint.2011.06.006.
  76. ^ a b Mol, D.; Shoshani, J.; Tikhonov, A.; van Geel, B.; Sano, S.; Lasarev, P.; Agenbroad, L. (2006). "The Yukagir mammoth: brief history, 14c dates, individual age, gender, size, physical and environmental conditions and storage". Scientific Annals, School of Geology Aristotle University of Thessaloniki. 98: 299–314.
  77. ^ Lister, 2007. pp. 102–103
  78. ^ Cherney, Michael D.; Fisher, Daniel C.; Auchus, Richard J.; Rountrey, Adam N.; Selcer, Perrin; Shirley, Ethan A.; Beld, Scott G.; Buigues, Bernard; Mol, Dick; Boeskorov, Gennady G.; Vartanyan, Sergey L.; Tikhonov, Alexei N. (2023). "Testosterone histories from tusks reveal woolly mammoth musth episodes". Nature. 617 (7961): 533–539. Bibcode:2023Natur.617..533C. doi:10.1038/s41586-023-06020-9. PMID 37138076. S2CID 258485513.
  79. ^ Lister, 2007. pp. 108–111
  80. ^ Rothschild, Bruce M.; Wang, Xiaoming; Shoshani, Jeheskel (September 1994). "Spondyloarthropathy in Proboscideans". Journal of Zoo and Wildlife Medicine. 25 (3): 360–366. JSTOR 20095389. Retrieved 17 April 2024.
  81. ^ Reumer, J. W. F.; Ten Broek, C. M. A.; Galis, F. (2014). "Extraordinary incidence of cervical ribs indicates vulnerable condition in Late Pleistocene mammoths". PeerJ. 2: e318. doi:10.7717/peerj.318. PMC 3970796. PMID 24711969.
  82. ^ Haynes, Gary; Klimowicz, Janis (27 August 2015). "A preliminary review of bone and teeth abnormalities seen in recent Loxodonta and extinct Mammuthus and Mammut, and suggested implications". Quaternary International. 379: 135–146. doi:10.1016/j.quaint.2015.04.001. Retrieved 19 April 2024 – via Elsevier Science Direct.
  83. ^ Lister, 2007. p. 87
  84. ^ a b c Lister, 2007. pp. 88–89
  85. ^ Lister, 2007. pp. 108–109
  86. ^ Pavelková Řičánková, V.; Robovský, J.; Riegert, J. (13 January 2014). "Ecological Structure of Recent and Last Glacial Mammalian Faunas in Northern Eurasia: The Case of Altai-Sayan Refugium". PLOS ONE. 9 (1): e85056. Bibcode:2014PLoSO...985056P. doi:10.1371/journal.pone.0085056. PMC 3890305. PMID 24454791.
  87. ^ Willerslev, E.; Davison, J.; Moora, M.; Zobel, M.; Coissac, E.; Edwards, M. E.; Lorenzen, E. D.; Vestergård, M.; Gussarova, G.; Haile, J.; Craine, J.; Gielly, L.; Boessenkool, S.; Epp, L. S.; Pearman, P. B.; Cheddadi, R.; Murray, D.; Bråthen, K. A.; Yoccoz, N.; Binney, H.; Cruaud, C.; Wincker, P.; Goslar, T.; Alsos, I. G.; Bellemain, E.; Brysting, A. K.; Elven, R.; Sønstebø, J. R. H.; Murton, J.; et al. (2014). "Fifty thousand years of Arctic vegetation and megafaunal diet" (PDF). Nature. 506 (7486): 47–51. Bibcode:2014Natur.506...47W. doi:10.1038/nature12921. PMID 24499916. S2CID 4461741.
  88. ^ Takahashi, K.; Wei, G.; Uno, H.; Yoneda, M.; Jin, C.; Sun, C.; Zhang, S.; Zhong, B. (2007). "AMS 14C chronology of the world's southernmost woolly mammoth (Mammuthus primigenius Blum.)". Quaternary Science Reviews. 26 (7–8): 954–957. Bibcode:2007QSRv...26..954T. doi:10.1016/j.quascirev.2006.12.001.
  89. ^ Álvarez-Lao, D. J.; García, N. (2012). "Comparative revision of the Iberian woolly mammoth (Mammuthus primigenius) record into a European context". Quaternary Science Reviews. 32: 64–74. Bibcode:2012QSRv...32...64A. doi:10.1016/j.quascirev.2011.11.004.
  90. ^ Diego J. Alvarez-Lao; et al. (2009), "The Padul mammoth finds — On the southernmost record of Mammuthus primigenius in Europe and its southern spread during the Late Pleistocene" (PDF), Palaeogeography, Palaeoclimatology, Palaeoecology, 278 (1–4): 57–70, Bibcode:2009PPP...278...57A, doi:10.1016/j.palaeo.2009.04.011
  91. ^ Gilbert, M. T. P.; Drautz, D. I.; Lesk, A. M.; Ho, S. Y. W.; Qi, J.; Ratan, A.; Hsu, C. -H.; Sher, A.; Dalen, L.; Gotherstrom, A.; Tomsho, L. P.; Rendulic, S.; Packard, M.; Campos, P. F.; Kuznetsova, T. V.; Shidlovskiy, F.; Tikhonov, A.; Willerslev, E.; Iacumin, P.; Buigues, B.; Ericson, P. G. P.; Germonpre, M.; Kosintsev, P.; Nikolaev, V.; Nowak-Kemp, M.; Knight, J. R.; Irzyk, G. P.; Perbost, C. S.; Fredrikson, K. M.; Harkins, T. T. (2008). "Intraspecific phylogenetic analysis of Siberian woolly mammoths using complete mitochondrial genomes". Proceedings of the National Academy of Sciences. 105 (24): 8327–8332. Bibcode:2008PNAS..105.8327G. doi:10.1073/pnas.0802315105. PMC 2423413. PMID 18541911.
  92. ^ Szpak, P.; Gröcke, D. R.; Debruyne, R.; MacPhee, R. D. E.; Guthrie, R. D.; Froese, D.; Zazula, G. D.; Patterson, W. P.; Poinar, H. N. (2010). "Regional differences in bone collagen δ13C and δ15N of Pleistocene mammoths: Implications for paleoecology of the mammoth steppe". Palaeogeography, Palaeoclimatology, Palaeoecology. 286 (1–2): 88–96. Bibcode:2010PPP...286...88S. doi:10.1016/j.palaeo.2009.12.009.
  93. ^ Stuart, A. J. (2005). "The extinction of woolly mammoth (Mammuthus primigenius) and straight-tusked elephant (Palaeoloxodon antiquus) in Europe" (PDF). Quaternary International. 126–128: 171–177. Bibcode:2005QuInt.126..171S. doi:10.1016/j.quaint.2004.04.021.
  94. ^ Debruyne, R.; Chu, G.; King, C. E.; Bos, K.; Kuch, M.; Schwarz, C.; Szpak, P.; Gröcke, D. R.; Matheus, P.; Zazula, G.; Guthrie, D.; Froese, D.; Buigues, B.; De Marliave, C.; Flemming, C.; Poinar, D.; Fisher, D.; Southon, J.; Tikhonov, A. N.; MacPhee, R. D. E.; Poinar, H. N. (2008). "Out of America: Ancient DNA Evidence for a New World Origin of Late Quaternary Woolly Mammoths". Current Biology. 18 (17): 1320–1326. doi:10.1016/j.cub.2008.07.061. PMID 18771918. S2CID 18663366.
  95. ^ Lister, 2007. pp. 116–117
  96. ^ a b Lister, 2007. pp. 118–125
  97. ^ a b Braun, I. M.; Palombo, M. R. (2012). "Mammuthus primigenius in the cave and portable art: An overview with a short account on the elephant fossil record in Southern Europe during the last glacial". Quaternary International. 276–277: 61–76. Bibcode:2012QuInt.276...61B. doi:10.1016/j.quaint.2012.07.010.
  98. ^ Demay, L.; Péan, S.; Patou-Mathis, M. (October 2012). "Mammoths used as food and building resources by Neanderthals: Zooarchaeological study applied to layer 4, Molodova I (Ukraine)" (PDF). Quaternary International. 276–277: 212–226. Bibcode:2012QuInt.276..212D. doi:10.1016/j.quaint.2011.11.019. hdl:2268/190618.
  99. ^ Lister, 2007. pp. 128–132
  100. ^ Lister, 2007. pp. 131–137
  101. ^ Lister, 2007. pp. 151–155
  102. ^ Wojtal, Piotr (2019). "The earliest direct evidence of mammoth hunting in Central Europe". Quaternary Science Reviews. 213: 162–166. doi:10.1016/j.quascirev.2019.04.004. S2CID 149647112.
  103. ^ Mussi, M.; Villa, P. (2008). "Single carcass of Mammuthus primigenius with lithic artifacts in the Upper Pleistocene of northern Italy" (PDF). Journal of Archaeological Science. 35 (9): 2606–2613. Bibcode:2008JArSc..35.2606M. doi:10.1016/j.jas.2008.04.014.
  104. ^ a b Aviss, B. (4 April 2012). "Woolly mammoth carcass may have been cut into by humans". BBC. Archived from the original on 6 April 2012. Retrieved 9 April 2012.
  105. ^ Nikolskiy, P.; Pitulko, V. (2013). "Evidence from the Yana Palaeolithic site, Arctic Siberia, yields clues to the riddle of mammoth hunting". Journal of Archaeological Science. 40 (12): 4189–4197. Bibcode:2013JArSc..40.4189N. doi:10.1016/j.jas.2013.05.020.
  106. ^ Overstreet, D. F.; Kolb, M. F. (2003). "Geoarchaeological contexts for Late Pleistocene archaeological sites with human-modified woolly mammoth remains in southeastern Wisconsin, U.S.A". Geoarchaeology. 18: 91–114. doi:10.1002/gea.10052. S2CID 129431648.
  107. ^ Joyce, D. J. (2006). "Chronology and new research on the Schaefer mammoth (?Mammuthus primigenius) site, Kenosha County, Wisconsin, USA". Quaternary International. 142–143: 44–57. Bibcode:2006QuInt.142...44J. doi:10.1016/j.quaint.2005.03.004.
  108. ^ a b Murchie, Tyler J.; Monteath, Alistair J.; Mahony, Matthew E.; Long, George S.; Cocker, Scott; Sadoway, Tara; Karpinski, Emil; Zazula, Grant; MacPhee, Ross D. E.; Froese, Duane; Poinar, Hendrik N. (2021). "Collapse of the mammoth-steppe in central Yukon as revealed by ancient environmental DNA". Nature Communications. 12 (7120 (2021)): 2031. Bibcode:2007QSRv...26.2031B. doi:10.1038/s41467-021-27439-6. PMC 8654998. PMID 34880234.
  109. ^ Nogués-Bravo, David; Rodríguez, Jesús; Hortal, Joaquín; Batra, Persaram; Araújo, Miguel B. (1 April 2008). "Climate Change, Humans, and the Extinction of the Woolly Mammoth". PLOS Biology. 6 (4): e79. doi:10.1371/journal.pbio.0060079. ISSN 1545-7885. PMC 2276529. PMID 18384234.
  110. ^ Stuart, Anthony J.; Lister, Adrian M. (2007). "Patterns of Late Quaternary megafaunal extinctions in Europe and northern Asia". CFS Courier Forschungsinstitut Senckenberg. 259: 289–299.
  111. ^ a b Lister, 2007. pp. 146–148
  112. ^ Palkopoulou, Eleftheria; Dalén, Love; Lister, Adrian M.; Vartanyan, Sergey; Sablin, Mikhail; Sher, Andrei; Edmark, Veronica Nyström; Brandström, Mikael D.; Germonpré, Mietje; Barnes, Ian; Thomas, Jessica A. (2013). "Holarctic genetic structure and range dynamics in the woolly mammoth". Proceedings of the Royal Society B. 280 (1770): 1–10. doi:10.1098/rspb.2013.1910. PMC 3779339. PMID 24026825.
  113. ^ Kahlke, Ralk-Dietrich (2015). "The maximum geographic extension of Late Pleistocene Mammuthus primigenius (Proboscidea, Mammalia) and its limiting factors". Quaternary International. 379: 147–154. Bibcode:2015QuInt.379..147K. doi:10.1016/j.quaint.2015.03.023.
  114. ^ a b Dehasque, Marianne; Pečnerová, Patrícia; Muller, Héloïse; Tikhonov, Alexei; Nikolskiy, Pavel; Tsigankova, Valeriya I.; Danilov, Gleb K.; Díez-del-Molino, David; Vartanyan, Sergey; Dalén, Love; Lister, Adrian M. (May 2021). "Combining Bayesian age models and genetics to investigate population dynamics and extinction of the last mammoths in northern Siberia". Quaternary Science Reviews. 259: 106913. Bibcode:2021QSRv..25906913D. doi:10.1016/j.quascirev.2021.106913.
  115. ^ Murchie, Tyler J.; Monteath, Alistair J.; Mahony, Matthew E.; Long, George S.; Cocker, Scott; Sadoway, Tara; Karpinski, Emil; Zazula, Grant; MacPhee, Ross D. E.; Froese, Duane; Poinar, Hendrik N. (8 December 2021). "Collapse of the mammoth-steppe in central Yukon as revealed by ancient environmental DNA". Nature Communications. 12 (1): 7120. Bibcode:2021NatCo..12.7120M. doi:10.1038/s41467-021-27439-6. ISSN 2041-1723. PMC 8654998. PMID 34880234.
  116. ^ a b Fordham, Damien A.; Brown, Stuart C.; Akçakaya, H. Reşit; Brook, Barry W.; Haythorne, Sean; Manica, Andrea; Shoemaker, Kevin T.; Austin, Jeremy J.; Blonder, Benjamin; Pilowsky, Julia; Rahbek, Carsten (2022). "Process-explicit models reveal pathway to extinction for woolly mammoth using pattern-oriented validation". Ecology Letters. 25 (1): 125–137. doi:10.1111/ele.13911. hdl:11343/299174. ISSN 1461-0248. PMID 34738712. S2CID 243762825.
  117. ^ a b Stuart, A. J.; Sulerzhitsky, L. D.; Orlova, L. A.; Kuzmin, Y. V.; Lister, A. M. (2002). "The latest woolly mammoths (Mammuthus primigenius Blumenbach) in Europe and Asia: A review of the current evidence" (PDF). Quaternary Science Reviews. 21 (14–15): 1559–1569. Bibcode:2002QSRv...21.1559S. doi:10.1016/S0277-3791(02)00026-4.
  118. ^ Lister, Adrian M. (18 June 2009). "Late‐glacial mammoth skeletons ( Mammuthus primigenius ) from Condover (Shropshire, UK): anatomy, pathology, taphonomy and chronological significance". Geological Journal. 44 (4): 447–479. doi:10.1002/gj.1162. ISSN 0072-1050. Retrieved 10 June 2024 – via Wiley Online Library.
  119. ^ Dale Guthrie, R. (2004). "Radiocarbon evidence of mid-Holocene mammoths stranded on an Alaskan Bering Sea island". Nature. 429 (6993): 746–749. Bibcode:2004Natur.429..746D. doi:10.1038/nature02612. PMID 15201907. S2CID 186242235.
  120. ^ Yesner, D. R.; Veltre, D. W.; Crossen, K. J.; Graham, R. W. "5,700-year-old Mammoth Remains from Qagnax Cave, Pribilof Islands, Alaska". Second World of Elephants Congress, (Hot Springs: Mammoth Site, 2005): 200–203.
  121. ^ Crossen, K. S. (2005). "5,700-Year-Old Mammoth Remains from the Pribilof Islands, Alaska: Last Outpost of North America Megafauna". Geological Society of America. 37: 463. Archived from the original on 3 March 2016. Retrieved 13 February 2020.
  122. ^ Graham, R. W.; Belmecheri, S.; Choy, K.; Culleton, B. J.; Davies, L. J.; Froese, D.; Heintzman, P. D.; Hritz, C.; Kapp, J. D.; Newsom, L. A.; Rawcliffe, R.; Saulnier-Talbot, É.; Shapiro, B.; Wang, Y.; Williams, J. W.; Wooller, M. J. (1 August 2016). "Timing and causes of mid-Holocene mammoth extinction on St. Paul Island, Alaska". Proceedings of the National Academy of Sciences. 113 (33): 9310–4. Bibcode:2016PNAS..113.9310G. doi:10.1073/pnas.1604903113. PMC 4995940. PMID 27482085.
  123. ^ Markus Milligan. "Mammoths still walked the earth when the Great Pyramid was being built". HeritageDaily – Heritage & Archaeology News. Archived from the original on 30 June 2015. Retrieved 5 July 2015.
  124. ^ Stuart, A. J.; Kosintsev, P. A.; Higham, T. F. G.; Lister, A. M. (2004). "Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth" (PDF). Nature. 431 (7009): 684–689. Bibcode:2004Natur.431..684S. doi:10.1038/nature02890. PMID 15470427. S2CID 4415073.
  125. ^ Vartanyan, S. L.; et al. (1995). "Radiocarbon Dating Evidence for Mammoths on Wrangel Island, Arctic Ocean, until 2000 BC". Radiocarbon. 37 (1): 1–6. Bibcode:1995Radcb..37....1V. doi:10.1017/S0033822200014703. ISSN 0033-8222. Archived from the original on 2 April 2012.
  126. ^ a b c d e Nyström, V.; Humphrey, J.; Skoglund, P.; McKeown, N. J.; Vartanyan, S.; Shaw, P. W.; Lidén, K.; Jakobsson, M.; Barnes, I. A. N.; Angerbjörn, A.; Lister, A.; Dalén, L. (2012). "Microsatellite genotyping reveals end-Pleistocene decline in mammoth autosomal genetic variation". Molecular Ecology. 21 (14): 3391–3402. doi:10.1111/j.1365-294X.2012.05525.x. PMID 22443459.
  127. ^ Wang, Y; Pedersen, M.W.; Alsos, I.g.; et al. (2021). "Late Quaternary dynamics of Arctic biota from ancient environmental genomics". Nature. 600 (7887): 86–92. Bibcode:2021Natur.600...86W. doi:10.1038/s41586-021-04016-x. PMC 8636272. PMID 34671161.
  128. ^ Seeber, Pa; Batke, L; Dvornikov, Y; Schmidt, A; Wang, Y; Stoof-Leichsenring, Kr; Moon, Kl; Shapiro, B; Epp, Ls (1 September 2023). Mitochondrial genomes of Pleistocene megafauna retrieved from recent sediment layers of two Siberian lakes (Report). elife. doi:10.7554/elife.89992.1.
  129. ^ a b Palkopoulou, E.; Mallick, S.; Skoglund, P.; Enk, J.; Rohland, N.; Li, H.; Omrak, A.; Vartanyan, S.; Poinar, H.; Götherström, A.; Reich, D.; Dalén, L. (23 April 2015). "Complete Genomes Reveal Signatures of Demographic and Genetic Declines in the Woolly Mammoth". Current Biology. 25 (10): 1395–1400. doi:10.1016/j.cub.2015.04.007. PMC 4439331. PMID 25913407.
  130. ^ Dunham, W. (24 April 2015). "Lonely end for the world's last woolly mammoths". ABC Science. Reuters. Archived from the original on 26 April 2015. Retrieved 24 April 2015.
  131. ^ Nystrom, V.; Dalen, L.; Vartanyan, S.; Liden, K.; Ryman, N.; Angerbjorn, A. (2010). "Temporal genetic change in the last remaining population of woolly mammoth". Proceedings of the Royal Society B: Biological Sciences. 277 (1692): 2331–2337. doi:10.1098/rspb.2010.0301. PMC 2894910. PMID 20356891.
  132. ^ a b Barsh, G. S.; Rogers, R. L.; Slatkin, M. (2017). "Excess of genomic defects in a woolly mammoth on Wrangel Island". PLOS Genetics. 13 (3): e1006601. doi:10.1371/journal.pgen.1006601. PMC 5333797. PMID 28253255.
  133. ^ Switek, B. (2017). "Dying woolly mammoths were in 'genetic meltdown'". Nature. doi:10.1038/nature.2017.21575. S2CID 184732688.
  134. ^ Arppe, L.; Karhu, J. A.; Vartanyan, S.; Drucker, D. G.; Etu-Sihvola, H.; Bocherens, H. (2019). "Thriving or surviving? The isotopic record of the Wrangel Island woolly mammoth population". Quaternary Science Reviews. 222: 105884. Bibcode:2019QSRv..22205884A. doi:10.1016/j.quascirev.2019.105884.
  135. ^ Ackerman, R. E. (1998). "Early maritime traditions in the Bering, Chukchi, and East Siberian seas". Arctic Anthropology. 35 (1): 247–262. JSTOR 40316468.
  136. ^ a b Fiedel, Stuart (2009). "Sudden Deaths: The Chronology of Terminal Pleistocene Megafaunal Extinction". In Haynes, G. (ed.). American Megafaunal Extinctions at the End of the Pleistocene. Vertebrate Paleobiology and Paleoanthropology. Springer. pp. 21–37. doi:10.1007/978-1-4020-8793-6_2. ISBN 978-1-4020-8792-9.
  137. ^ Graham, R. W.; Belmecheri, S.; Choy, K.; Culleton, B. J.; Davies, L. J.; Froese, D.; Heintzman, P. D.; Hritz, C.; Kapp, J. D.; Newsom, L. A.; Rawcliffe, R.; Saulnier-Talbot, É.; Shapiro, B.; Wang, Y.; Williams, J. W.; Wooller, M. J. (1 August 2016). "Timing and causes of mid-Holocene mammoth extinction on St. Paul Island, Alaska". Proceedings of the National Academy of Sciences. 113 (33): 9310–4. Bibcode:2016PNAS..113.9310G. doi:10.1073/pnas.1604903113. PMC 4995940. PMID 27482085.
  138. ^ Nogués-Bravo, D.; Rodríguez, J. S.; Hortal, J. N.; Batra, P.; Araújo, M. B. (2008). Barnosky, Anthony (ed.). "Climate Change, Humans, and the Extinction of the Woolly Mammoth". PLOS Biology. 6 (4): e79. doi:10.1371/journal.pbio.0060079. PMC 2276529. PMID 18384234.
  139. ^ Sedwick, C. (2008). "What Killed the Woolly Mammoth?". PLOS Biology. 6 (4): e99. doi:10.1371/journal.pbio.0060099. PMC 2276526. PMID 20076709.
  140. ^ Martin, P. S (2005). Twilight of the Mammoths: Ice Age Extinctions and the Rewilding of America. University of California Press. pp. 165–173. ISBN 978-0-520-23141-2.
  141. ^ Burney, D.; Flannery, T. (2005). "Fifty millennia of catastrophic extinctions after human contact" (PDF). Trends in Ecology & Evolution. 20 (7): 395–401. doi:10.1016/j.tree.2005.04.022. PMID 16701402. Archived from the original (PDF) on 10 June 2010.
  142. ^ Doughty, C. E.; Wolf, A.; Field, C. B. (2010). "Biophysical feedbacks between the Pleistocene megafauna extinction and climate: the first human-induced global warming?". Geophysical Research Letters. 37 (15): L15703. Bibcode:2010GeoRL..3715703D. doi:10.1029/2010GL043985.
  143. ^ a b c Lister, 2007. pp. 45–75
  144. ^ a b Lister, 2007. pp. 50–53
  145. ^ a b Pfizenmayer, E. W. (1939). Siberian Man and Mammoth. London: Blackie and Son. pp. 46–61.
  146. ^ Vereshchagin, N. K. (2009). "The mammoth "cemeteries" of north-east Siberia". Polar Record. 17 (106): 3–12. doi:10.1017/S0032247400031296. S2CID 129654739.
  147. ^ a b c Tolmachoff, I. P. (1929). "The carcasses of the mammoth and rhinoceros found in the frozen ground of Siberia". Transactions of the American Philosophical Society. 23 (1): 11–23. doi:10.2307/1005437. JSTOR 1005437.
  148. ^ Adams, M. (1808). "Some Account of a Journey to the Frozen-Sea, and of the Discovery of the Remains of a Mammoth". The Philadelphia Medical and Physical Journal. 3: 120–137.
  149. ^ Tilesio, W. G. (1815). "De skeleto mammonteo Sibirico ad maris glacialis littora anno 1807 effosso, cui praemissae Elephantini generis specierum distinctiones". Mémoires de l'Académie Impériale des Sciences de St. Pétersbourg (in Latin). 5: 406–514.
  150. ^ Cohen, C. (2002). The Fate of the Mammoth: Fossils, Myth, and History. University of Chicago Press. p. 113. ISBN 978-0-226-11292-3.
  151. ^ Pfizenmayer, E. (1907). "A Contribution to the Morphology of the Mammoth, Elephas Primigenius Blumenbach; With an Explanation of My Attempt at a Restoration". Annual Report of the Board of Regents of the Smithsonian Institution: 326–334.
  152. ^ Farrand, W. R. (1961). "Frozen Mammoths and Modern Geology: The death of the giants can be explained as a hazard of tundra life, without evoking catastrophic events". Science. 133 (3455): 729–735. Bibcode:1961Sci...133..729F. doi:10.1126/science.133.3455.729. PMID 17777646.
  153. ^ Lister, 2007. pp. 57–58
  154. ^ Mol, D. et al. (2001). "The Jarkov Mammoth: 20,000-Year-Old carcass of a Siberian woolly mammoth Mammuthus primigenius (Blumenbach, 1799)". The World of Elephants, Proceedings of the 1st International Congress ( 16–20 October 2001, Rome): 305–309. Full text pdf
  155. ^ Debruyne, R. G.; Barriel, V. R.; Tassy, P. (2003). "Mitochondrial cytochrome b of the Lyakhov mammoth (Proboscidea, Mammalia): New data and phylogenetic analyses of Elephantidae". Molecular Phylogenetics and Evolution. 26 (3): 421–434. doi:10.1016/S1055-7903(02)00292-0. PMID 12644401.
  156. ^ Kosintsev, P. A.; Lapteva, E. G.; Trofimova, S. S.; Zanina, O. G.; Tikhonov, A. N.; Van Der Plicht, J. (2012). "Environmental reconstruction inferred from the intestinal contents of the Yamal baby mammoth Lyuba (Mammuthus primigenius Blumenbach, 1799)" (PDF). Quaternary International. 255: 231–238. Bibcode:2012QuInt.255..231K. doi:10.1016/j.quaint.2011.03.027. S2CID 129303118.
  157. ^ Rincon, P. (10 July 2007). "Baby mammoth discovery unveiled". BBC News. Archived from the original on 11 August 2007. Retrieved 13 July 2007.
  158. ^ Solovyov, D. (11 July 2007). "Baby mammoth find promises breakthrough". Reuters. Archived from the original on 13 July 2007. Retrieved 13 July 2007.
  159. ^ Smith, O. (21 April 2009). "Baby mammoth Lyuba, pristinely preserved, offers scientists rare look into mysteries of Ice Age". Daily News. New York. Archived from the original on 15 August 2009.
  160. ^ Fisher, Daniel C. (2014). "X-ray computed tomography of two mammoth calf mummies". Journal of Paleontology. 88 (4): 664–675. Bibcode:2014JPal...88..664F. doi:10.1666/13-092. S2CID 28393815.
  161. ^ Mashchenko, E. N.; Protopopov, A. V.; Plotnikov, V. V.; Pavlov, I. S. (2013). "Specific characters of the mammoth calf (Mammuthus primigenius) from the Khroma River (Yakutia)". Biology Bulletin. 40 (7): 626–641. doi:10.1134/S1062359013070042. S2CID 16675371.
  162. ^ Yamagata, K.; Nagai, K.; Miyamoto, H.; Anzai, M.; Kato, H.; Miyamoto, K.; Kurosaka, S.; Azuma, R.; Kolodeznikov, I. I.; Protopopov, A. V.; Plotnikov, V. V.; Kobayashi, H.; Kawahara-Miki, R.; Kono, T.; Uchida, M.; Shibata, Y.; Handa, T.; Kimura, H.; Hosoi, Y.; Mitani, T.; Matsumoto, K.; Iritani, A. (2019). "Signs of biological activities of 28,000-year-old mammoth nuclei in mouse oocytes visualized by live-cell imaging". Scientific Reports. 9 (1): 4050. Bibcode:2019NatSR...9.4050Y. doi:10.1038/s41598-019-40546-1. PMC 6411884. PMID 30858410.
  163. ^ Wong, K. (2013). "Can a mammoth carcass really preserve flowing blood and possibly live cells?". Nature. doi:10.1038/nature.2013.13103. S2CID 87298066.
  164. ^ Reardon, Sophie (2022). "Rare mummified baby woolly mammoth with skin and hair found in Canada". Retrieved 26 June 2022.
  165. ^ Proulx, Michel (24 June 2022). "'She's perfect and she's beautiful': Frozen baby woolly mammoth discovered in Yukon gold fields". Canadian Broadcasting Corporation. Retrieved 8 November 2022.
  166. ^ Bringing them Back to Life Archived 2017-03-29 at the Wayback Machine. Carl Zimmer, National Geographic. April 2013.
  167. ^ a b c Lister, 2007. pp. 42–43
  168. ^ Ghosh, Pallab (23 April 2015). "Mammoth genome sequence completed". BBC News. Archived from the original on 24 April 2015.
  169. ^ The Long Now Foundation – Revive and Restore Archived 24 April 2015 at the Wayback Machine.
  170. ^ Can scientists bring mammoths back to life by cloning? Archived 8 October 2017 at the Wayback Machine Jackson Landers. 9 February 2015. The Washington Post.
  171. ^ Webster, Ben (23 March 2015). "Scientist takes mammoth-cloning a step closer". The Sunday Times.
  172. ^ Sarah Fecht (24 March 2014), Woolly Mammoth DNA Successfully Spliced Into Elephant Cells, Popular Science, archived from the original on 26 March 2015
  173. ^ Zimov, S. A. (2005). "Essays on Science and Society: Pleistocene Park: Return of the Mammoth's Ecosystem". Science. 308 (5723): 796–798. doi:10.1126/science.1113442. PMID 15879196.
  174. ^ Rohwer, Y.; Marris, E. (2018). "An analysis of potential ethical justifications for mammoth de-extinction and a call for empirical research". Ethics, Policy & Environment. 21 (1): 127–142. doi:10.1080/21550085.2018.1448043. S2CID 158056898.
  175. ^ Griffin, A. (23 March 2015). "Woolly mammoth could be revived after scientists paste DNA into elephant's genetic code". The Independent. Archived from the original on 25 September 2015.
  176. ^ Loi, Pasqualino; Saragusty, Joseph; Ptak, Grazyna (2014). "Cloning the Mammoth: A Complicated Task or Just a Dream?". Reproductive Sciences in Animal Conservation. Advances in Experimental Medicine and Biology. Vol. 753. pp. 489–502. doi:10.1007/978-1-4939-0820-2_19. ISBN 978-1-4939-0819-6. PMID 25091921.
  177. ^ "Woolly mammoths are being brought back from extinction by scientists". euronews. 17 September 2021. Retrieved 19 September 2021.
  178. ^ Carlson, Kara. "Could Austin entrepreneur's company help bring back the woolly mammoth?". Austin American-Statesman. Retrieved 19 September 2021.
  179. ^ Newcomb, Raymond Lee (1888). Our lost explorers : the narrative of the Jeannette Arctic Expedition as related by the survivors, and in the records and last journals of Lieutenant De Long Archived 17 March 2016 at the Wayback Machine. p. 96
  180. ^ Patkanov, S. (1897), Die lrtysch-Ostjaken und ihre Volkspoesie, vol. I, St. Petersburg: St. Petersburg, pp. 123–124, archived from the original on 7 November 2018
  181. ^ Laufer, Berthold (1913), "Arabic and Chinese Trade in Walrus and Narwhal Ivory", T'oung Pao, Second Series, 14 (3): 329, doi:10.1163/156853213X00213, hdl:2027/hvd.32044009725912, JSTOR 4526349
  182. ^ Cohen, C. (2002). The Fate of the Mammoth: Fossils, Myth, and History. University of Chicago Press. pp. 197–198. ISBN 978-0-226-11292-3. Retrieved 10 August 2015.
  183. ^ a b Strong, W. D. (1934). "North American Indian traditions suggesting a knowledge of the mammoth". American Anthropologist. 36: 81–88. doi:10.1525/aa.1934.36.1.02a00060.
  184. ^ Lankford, G. E. (1980). "Pleistocene Animals in Folk Memory". The Journal of American Folklore. 93 (369): 294–296. doi:10.2307/540573. JSTOR 540573. {subscription required}
  185. ^ a b Mayor, A. (2005). Fossil Legends of the First Americans. Princeton: Princeton University Press. p. 97. ISBN 978-0-691-11345-6.
  186. ^ Lankford, G. E. (1980). "Pleistocene Animals in Folk Memory". The Journal of American Folklore. 93 (369): 293–304. doi:10.2307/540573. JSTOR 540573. {subscription required}
  187. ^ a b Lister, 2007. pp. 137–139
  188. ^ Larmer, B. (April 2013). "Mammoth Tusk Hunters". Archived from the original on 2 April 2013.
  189. ^ Lister, 2007. p. 54
  190. ^ Glass, J. R.; Davis, M.; Walsh, T. J.; Sargis, E. J.; Caccone, A.; Fiorillo, A. (2016). "Was Frozen Mammoth or Giant Ground Sloth Served for Dinner at The Explorers Club?". PLOS ONE. 11 (2): e0146825. Bibcode:2016PLoSO..1146825G. doi:10.1371/journal.pone.0146825. PMC 4740485. PMID 26840445.
  191. ^ Weiyun, T. (2011). "'Lucky Hands' in pursuit of dinosaurs". SHINE. Retrieved 4 July 2019.
  192. ^ Chun, Alex (30 March 2023). "This massive meatball was made With woolly mammoth DNA". Smithsonian Magazine. Retrieved 24 March 2024.
  193. ^ a b c Sjögren, B. (1962). Farliga djur och djur som inte finns (in Swedish). Prisma. p. 168.
  194. ^ Lister, 2007. p. 55
  195. ^ Murray, M. (1960). "Henry Tukeman: Mammoth's Roar was Heard All The Way to the Smithsonian". Tacoma Public Library. Archived from the original on 18 January 2012. Retrieved 17 January 2008.
  196. ^ McCarthy, Michael (28 September 2009), The Big Question: Are so-called 'extinct' species really extinct, and will we rediscover any?, retrieved 5 August 2019