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119-487: Stegoloxodon is an extinct genus of dwarf elephant known from the Early Pleistocene of Indonesia. It contains two species, S. indonesicus from Java , and S. celebensis from Sulawesi . Its relationship with other elephants is uncertain. S. celebensis was originally described as Archidiskodon celebensis in 1949 by Dirk Albert Hooijer . S. indonesicus was originally described by Miklós Kretzoi based on

238-459: 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. A 2011 study showed that light individuals would have been rare. A 2014 study instead indicated that the colouration of an individual varied from nonpigmented on

357-412: A 1975 publication to be found associated with lithic artefacts. The age of the find was considered to be uncertain, likely older than 9,000 years, but could not be dated precisely due to a lack of collagen. Additionally, an isolated tusk was reported from the northwest of the island. On Delos , an indeterminate dwarf elephant known from a third molar was reported in 1908. This specimen clearly belongs to

476-481: A body mass of around 1.7 tons, while the later Stegodon florensis insularis from the Late Pleistocene is estimated to be around 17% the size of mainland Stegodon species, with a shoulder height of around 130 cm (4.3 ft), and a body mass of about 570 kilograms (1,260 lb) Stegodon florensis became extinct about 50,000 years ago, around the time of the arrival of modern humans to Flores. During

595-571: A cast. 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

714-457: 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. 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

833-552: A dwarf species, but it is difficult to quantify its size precisely. On Naxos the species Palaeoloxodon lomolinoi has been described based on a partial skull including the maxilla bones and third molar teeth found near the Trypiti river, of probable Late Pleistocene age. It is estimated to be around 8% the size of P. antiquus , and had a smaller body size than that represented by the dwarf elephant from Delos. The Eastern Cyclades islands of Delos, Naxos , and Paros were connected during

952-430: 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. 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

1071-534: 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; 2.4–2.7 m (7 ft 10 in – 8 ft 10 in) and 45 kg (99 lb) was a more typical size. Female tusks were smaller and thinner, 1.5–1.8 m (4 ft 11 in – 5 ft 11 in) and weighing 9 kg (20 lb). For comparison,

1190-423: 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. 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

1309-528: 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 (5.9 in) long, and the sixth were about 30 cm (12 in) long and weighed 1.8 kg (4.0 lb). The molars grew larger and contained more ridges with each replacement. The woolly mammoth

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1428-412: A molar found near Bumiayu , originally attributed to Elephas planifrons , who coined the genus Stegoloxodon to contain the species. A 1973 paper argued for the synonymy of the two species, which was later rejected, though they are usually considered closely related. Some later papers referred the species to Elephas . A paper by Paul Yves Sondaar in 1984 started a trend of referring to the two species to

1547-402: A more extensive covering of hair than adults. This is thought to be for thermoregulation , helping them lose heat in their hot environments. Comparison between the over-hairs of woolly mammoths and extant elephants show that they did not differ much in overall morphology. Woolly mammoths had numerous sebaceous glands in their skin, which secreted oils into their hair; this would have improved

1666-423: A number other large mammal species, including herbivores and carnivores. The youngest records of this species on Sicily date to around 20,000 years ago, close to the time of arrival of modern humans on Sicily. The dwarf mammoth species Mammuthus lamarmorai descended from steppe mammoths ( Mammuthus trogontherii ) that colonised Sardinia sometime after 450,000 years ago. It is suggested to have survived into

1785-449: 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

1904-469: A result of reducing land area of the Japanese archipelago. The latest and smallest species S. aurorae is estimated to be 25% the size of its mainland ancestor with a body mass of around 2,122 kilograms (4,678 lb). During the late Middle Pleistocene to Late Pleistocene around 330,000-24,000 years ago, the Japanese archipelago was inhabited by the elephant species Palaeoloxodon naumanni . This species

2023-415: 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. Likewise, spondyloarthropathy has also been identified in woolly mammoth remains. An extra number of cervical vertebrae has been found in 33% of specimens from

2142-523: A shoulder height of up to 1.9 metres (6.2 ft), with a body mass of 630–890 kilograms (1,390–1,960 lb). Remains of the species are suggested to date to Late Pleistocene. Radiocarbon dating done in the 1970s suggested that the species survived until around 3,500 years ago, which would make the latest surviving Palaeoloxodon species and the youngest elephant in Europe, but these dates are tentative and await corroboration by other research. On Astypalaia ,

2261-472: A similar relationship with M. trogontherii in areas where their range overlapped. In 2021, DNA older than a million years was sequenced for the first time, from two mammoth teeth of Early Pleistocene age found in eastern Siberia. One tooth from Adycha (1–1.3 million years old) belonged to a lineage that was ancestral to later woolly mammoths, whereas the other from Krestovka (1.1–1.65 million years old) belonged to new lineage. The study found that half of

2380-503: A single tusk of a dwarf elephant of unknown age was excavated in the late 1990s. Due to the isolated status of the island it very likely represents an endemic species. Though the size of the animal is difficult to constrain precisely, it was probably similar in size to P. tiliensis . On Kasos , which during the Pleistocene was connected with the islands of Karpathos and Saria , a single dwarf Palaeoloxodon molar has been found. Due to

2499-471: A strong smell called temporin . Their fur may have helped in spreading the scent further. 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. 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

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2618-490: A succession of endemic dwarf species of Stegodon, probably representing a single lineage lived in the Japanese archipelago , probably derived from the mainland Chinese S. zydanskyi. In chronological succession these species are Stegodon miensis (4-3 million years ago) Stegodon protoaurorae (3-2 million years ago) and Stegodon aurorae , (2-1 million years ago) which show a progressive size reduction through time, possibly as

2737-520: 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 (0.98 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

2856-509: A tropical animal would be found in such a cold area as Siberia, and suggested that they might have been transported there by the Great Flood. In 1796, French biologist Georges Cuvier was the first to identify the woolly mammoth remains not as modern elephants transported to the Arctic, but as an entirely new species. He argued this species had gone extinct and no longer existed, a concept that

2975-433: A varied diet to support their growth, like modern elephants. An adult of 6 tonnes would need to eat 180 kg (400 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

3094-539: 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. Woolly mammoth dental enamel from Poland has demonstrated that woolly mammoths were seasonally migratory. Recurring shifts in δ O and Sr/ Sr found in layers of

3213-413: 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

3332-508: Is P. falconeri , which is one of the smallest dwarf elephant species at around 1 metre (3.3 ft) tall, and was strongly modified from its ancestor in numerous aspects, which lived in a depauperate fauna with no other large mammal species. Later, around 200,000 years ago, this species was replaced by a second colonisation by P. antiquus , which gave rise to the larger (though still considerably dwarfed) 2 metres (6.6 ft) tall species P. mnaidriensis , which on Sicily lived alongside

3451-407: Is also one of the smallest dwarf elephant species, comparable in size to P. falconeri , with an estimated shoulder height of 1 metre (3.3 ft). The species likely evolved from the earlier larger (though still strongly dwarfed) Palaeoloxodon xylophagou known from fossils dating to around 200,000 years ago. Remains of the species were first discovered and recorded by Dorothea Bate in a cave in

3570-410: Is considered to have had the most complex molars of any elephant. Adult woolly mammoths could effectively defend themselves from predators with their tusks, trunks and size; however, 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

3689-413: Is indicated on many preserved tusks by flat, polished sections up to 30 cm (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. The lifespan of mammals is related to their size. Since modern elephants can reach

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3808-520: Is not known outside of Africa) or to Elephas planifrons (which is known from the Late Pliocene-Early Pleistocene of South Asia). Both S. indonesicus and S. celebensis were dwarf elephants substantially smaller than mainland elephant species as a result of insular dwarfism , with S. celebensis estimated to have been around 150 cm (4.9 ft) tall. The molar plates of both species are low-crowned with thick enamel, with

3927-496: Is one the smallest known species, at around 8% of the size of its mainland ancestor, with an estimated body mass of around 250 kilograms (550 lb). The species Stegodon timorensis is known from the Middle Pleistocene of Timor. It is a small-sized species, only slightly larger than S. sondaarii, and around 23% the size of mainland species, with an estimated body mass of around 770 kilograms (1,700 lb). On Luzon

4046-423: 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

4165-477: Is short and tall, with fronto-parietal crests, and slightly curved upper tusks. Isotope analysis suggests that S. celebensis was a flexible feeder, while S. indonesicus is suggested to have been a dedicated browser on C3 vegetation in forested habitats. S. celebensis inhabited the island alongside the similarly sized dwarf Stegodon species Stegodon sompoensis, and a larger unnamed Stegodon species while Stegoxolodon indonesicus co-existed alongside

4284-546: Is the dwarf Middle-Late Pleistocene Sardinian mammoth , Mammuthus lamarmorai (Major, 1883), the first endemic elephant of the Mediterranean islands recognized as belonging to the mammoth line. Mammuthus creticus from the Early Pleistocene of Crete, formerly considered a member of Palaeoloxodon , is now also considered to be a mammoth, and approaches the size of the smallest dwarf elephants. During low sea levels,

4403-420: Is the size of the pelvic girdle, since the opening that functions as the birth canal is always wider in females than in males. Though the mammoths on Wrangel Island were smaller than those of the mainland, their size varied, and they were not small enough to be considered " island dwarfs ". The last woolly mammoth populations are claimed to have decreased in size and increased their sexual dimorphism , but this

4522-726: Is uncertain; depending on author, they are either considered primitive forms of an advanced species or advanced forms of a primitive species. Distinguishing and determining these intermediate forms has been called one of the most long-lasting and complicated problems in Quaternary palaeontology. Regional and intermediate species and subspecies such as M. intermedius , M. chosaricus , M. p. primigenius , M. p. jatzkovi , M. p. sibiricus , M. p. fraasi , M. p. leith-adamsi , M. p. hydruntinus , M. p. astensis , M. p. americanus , M. p. compressus , and M. p. alaskensis have been proposed. A 2011 genetic study showed that two examined specimens of

4641-769: The African bush elephant ( Loxodonta africana ), but is considerably smaller than the earlier Mammuthus meridionalis and Mammuthus trogontherii and the contemporary Mammuthus columbi . 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 m (9 ft 10 in – 10 ft 10 in) tall and 5.2–6.9 t (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 m (8 ft 9 in – 9 ft 8 in) tall and 3.9–5.2 t (8,600–11,500 lb) in weight). One of

4760-448: 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. The American president Thomas Jefferson , who had a keen interest in palaeontology , was partially responsible for transforming

4879-585: The Kyrenia hills of northern Cyprus in 1902 and reported in 1903. A population of the Columbian mammoth ( Mammuthus columbi ) arrived on the northern Channel Islands of California during the late Middle Pleistocene, around 250-150,000 years ago, giving rise to a dwarfed species, the pygmy mammoth ( Mammuthus exilis ). Channel Islands mammoths ranged from 150–190 cm (59–75 in) in shoulder height. These mammoths became extinct around 13,000 years ago, around

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4998-511: The Last Glacial Period , which suggests that the Delos species and P. lomolinoi were not contemporaneous, with the former possibly being the ancestor of the latter, though nothing can be said for certain. On Rhodes , bones of an unnamed endemic dwarf elephant have been discovered in cave deposits on the east coast. This elephant was similar in size to Palaeoloxodon mnaidriensis, around 20%

5117-617: The Middle Pleistocene . Woolly mammoths entered North America about 100,000 years ago by crossing the Bering Strait . 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

5236-637: The Proboscidea was published, wherein he used various taxon names that had previously been proposed for mammoth species, including replacing Mammuthus with Mammonteus , as he believed the former name to be invalidly published. Mammoth taxonomy was simplified by various researchers from the 1970s onwards, all species were retained in the genus Mammuthus , and many proposed differences between species were instead interpreted as intraspecific variation. Osborn chose two molars (found in Siberia and Osterode ) from Blumenbach's collection at Göttingen University as

5355-470: The lectotype specimens for the woolly mammoth, since holotype designation was not practised in Blumenbach's time, and one of the specimens Blumenbach had described had been of the unrelated straight-tusked elephant ( Palaeoloxodon antiquus ). Soviet palaeontologist Vera Gromova further proposed the former should be considered the lectotype with the latter as paralectotype. Both molars were thought lost by

5474-689: 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. Mammoth remains had long been known in Asia before they became known to Europeans. The origin of these remains was long a matter of debate and often explained as being remains of legendary creatures . The mammoth

5593-523: 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. 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

5712-585: The British physician 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 . 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. Others interpreted Sloane's conclusion slightly differently, arguing

5831-526: 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. 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

5950-492: The Cyclades archipelago. There are many uncertainties about the time of colonisation, the phylogenetic relationships and the taxonomic status of dwarf elephants on the Mediterranean islands. Extinction of the insular dwarf elephants has not been correlated with the arrival of humans to the islands. Furthermore, it has been suggested by the palaeontologist Othenio Abel in 1914, that the finding of skeletons of such elephants sparked

6069-485: The Last Glacial Period, until at least 60-30,000 years ago. Mammuthus creticus is known from remains probably dating to the Early Pleistocene . It likely descends from Mammuthus meridionalis . It is the smallest mammoth and is among the smallest dwarf elephants known, with a shoulder height of about 1 metre (3.3 ft) and a weight of about 180 kilograms (400 lb). Palaeoloxodon creutzburgi from

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6188-503: The Late Pliocene-Early Pleistocene on Sulawesi, two species of dwarf proboscidean coinhabited the island, the elephant Stegoloxodon celebensis , and Stegodon sompoensis. The former was about 150 cm (4.9 ft) tall, while the latter was around 32% the size of mainland Stegodon species, with an estimated body mass of about a ton. Later in the Pleistocene, these animals were replaced by larger-sized species of Stegodon and elephants, with an indeterminate Stegodon species from

6307-512: The Mediterranean islands during the Pleistocene , including all the major islands with the apparent exception of Corsica and the Balearics . Mediterranean dwarf elephants have generally been considered as members of the genus Palaeoloxodon , derived from the continental straight-tusked elephant , Palaeoloxodon antiquus ( Falconer & Cautley , 1847), Syn. : Elephas antiquus . An exception

6426-523: The Mediterranean islands were colonised again and again, giving rise, sometimes on the same island, to several species (or subspecies) of different body sizes. As the Ice Age came to an end, sea levels rose, stranding elephants on the island. The island of Sicily appears to have been colonised by proboscideans in at least three separate waves of colonisation. These endemic dwarf elephants were taxonomically different on each island or group of very close islands, like

6545-407: The Middle Pleistocene and Late Pleistocene is significantly larger, with an estimated body mass comparable to living Asian elephant, around 40% the size of its mainland ancestor. Remains of dwarf elephants have been briefly reported from Paros , Milos and Serifos in historical publications, but these lack any detailed information. On Kýthnos , the remains of a dwarf elephant were reported in

6664-566: The Middle Pleistocene of Sulawesi being around 57% the size of mainland species, with an estimated bodymass of about 2 tons. The species Stegodon trigonocephalus is known from the Early-Middle Pleistocene of Java. A population from the Trinil H.K locality, which likely dates to the Middle Pleistocene, is around 65% the size of mainland Stegodon species. Large individuals are estimated to have reached around 280 cm (9.2 ft) at

6783-412: 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

6902-588: 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

7021-401: 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. Woolly mammoths had four functional molar teeth at

7140-412: 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

7259-420: 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

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7378-468: The dwarf Stegodon luzonensis is known from remains found in the Manila Basin of an uncertain Pleistocene age, as well as remains found near the early Middle Pleistocene Nesorhinus butchery site dating to around 700,000 years ago. It is around 40% the size of mainland Stegodon species, with a body mass of around 1.3 tons. Though the temporal span of Stegodon on Luzon is not well constrained due to

7497-443: The earlier type, becoming the southern mammoth ( M. meridionalis ) about 2–1.7 million years ago. In turn, this species was replaced by the steppe mammoth ( M. trogontherii ) with 18–20 ridges, which evolved in eastern Asia around 1 million years ago. Mammoths derived from M. trogontherii evolved molars with 26 ridges 400,000 years ago in Siberia and became the woolly mammoth. The earliest identified forms of woolly mammoth date to

7616-519: The enamel correspond to seasonal variations and indicate that Polish woolly mammoths inhabited southern Poland during winter but grazed the Polish midlands during summer. 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

7735-431: The extreme polar variation in length of daylight. Similar mutations are known in other Arctic mammals, such as reindeer. A 2019 study of the woolly mammoth mitogenome suggest that these had metabolic adaptations related to extreme environments. 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

7854-503: 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. 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

7973-402: 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,

8092-420: 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

8211-522: 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. In 1738, the German zoologist Johann Philipp Breyne argued that mammoth fossils represented some kind of elephant. He could not explain why

8330-437: The genus " Elephas " with quotation remarks, reflecting uncertainty about their placement in the genus. A 2008 review by Markov and Saegusa resurrected the genus Stegoloxodon to house the two species. The relationship with other elephants has been considered uncertain, which has been suggested to be the result of dwarfism-caused changes to its morphology. Some authors have suggested a close relationship to Loxodonta (which

8449-777: The gomphothere Sinomastodon bumiajuensis , and an unnamed dwarf species of Stegodon . Dwarf elephant Fossil remains of dwarf elephants have been found on the Mediterranean islands of Cyprus , Malta , Crete , Sicily , Sardinia , the Cyclades Islands and the Dodecanese Islands , which are mostly members of the genus Palaeoloxodon , descending from the large 4 metres (13 ft) tall straight-tusked elephant ( Palaeoloxodon antiquus ) of mainland Europe , though two species represent dwarf mammoths . Dwarf species of elephants and Stegodon have been found on

8568-408: The hair follicles of both a 20,000-year-old mammoth retrieved from permafrost and another that died 60,000 years ago. In 2012, proteins were confidently identified for the first time, collected from a 43,000-year-old woolly mammoth. Since many remains of each species of mammoth are known from several localities, reconstructing the evolutionary history of the genus through morphological studies

8687-399: The hardest times for northern animals to survive. Examination of preserved calves shows that they were all born during spring and summer, and since modern elephants have gestation periods of 21–22 months, the mating season probably was from summer to autumn. δ15N isotopic analysis of the teeth of "Lyuba" has demonstrated their prenatal development , and indicates its gestation period

8806-425: The idea that they belonged to giant one-eyed monsters, because the center nasal opening was thought to be the socket of a single eye, and thus perhaps were, for example, the origin of the one-eyed Cyclopes of Greek mythology . Sicily and Malta were inhabited by two successive waves of dwarf elephants derived from P. antiquus, which first arrived on the islands at least 500,000 years ago. The first of these species

8925-678: 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. 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

9044-505: The islands of Indonesia and the Philippines, with dwarfed species of Stegodon also having been found in Japan. The Channel Islands of California once supported the pygmy mammoth , a dwarf species descended from Columbian mammoths , while the woolly mammoths that existed on Wrangel Island north of Siberia were once considered dwarfs, but are not anymore. Dwarf elephants first inhabited

9163-488: The lamellae frequency being notably higher in S. celebensis than in S. indonesicus . Both species retained permanent premolars . Lower tusks are present in some adult individuals of S. celebensis , unlike modern elephants, which is either suggested to have been a retained ancestral trait lost in modern elephants, or the result of paedomorphosis as a consequence of dwarfism, with the mandibular symphysis of S. celebensis being downwardly turned. The skull of S. celebensis

9282-438: 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,

9401-464: The largest recorded woolly mammoths is the Siegsdorf specimen from Germany, with an estimated shoulder height of 3.49 m (11.5 ft) and an estimated body mass of 8.2 t (18,000 lb). A newborn calf would have weighed about 90 kg (200 lb). Few frozen specimens have preserved genitals, so the sex is usually determined through examination of the skeleton. The best indication of sex

9520-427: The last glaciation 25–20,000 years ago show slower growth rates. 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

9639-506: 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

9758-418: The limited number of finds, remains are suggested to span from at least around 1-0.8 million years ago to around 400,000 years ago. The extinct dwarf elephant Elephas beyeri is also known from the island of an unknown (probably Pleistocene) age, which is estimated to have been about 1.2 metres (3.9 ft) in shoulder height. On the island of Mindanao , the dwarf Stegodon species Stegodon mindanensis

9877-440: 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 . Isotope analysis shows that woolly mammoths fed mainly on C3 plants , unlike horses and rhinos. Scientists identified milk in

9996-476: The mammoth and Asian elephant lineages diverged 5.8–7.8 million years ago, while African elephants diverged from an earlier common ancestor 6.6–8.8 million years ago. In 2008, much of the woolly mammoth's chromosomal DNA was mapped. The analysis showed that the woolly mammoth and the African elephant are 98.55% to 99.40% identical. The team mapped the woolly mammoth's nuclear genome sequence by extracting DNA from

10115-530: The mammoth genome project in 2015 sparked discussion about potentially reviving the woolly mammoth through several various methods. However, none of these approaches are currently feasible. Remains of woolly mammoths were long known by native Siberians and Native Americans , who had various ways of interpreting them. Remains later reached other parts of Asia and Europe, where they were also interpreted in various ways prior to modern science . The first woolly mammoth remains studied by scientists were examined by

10234-975: 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. The Asian elephant is the closest extant relative of the mammoths. The following cladogram shows the placement of the woolly mammoth among Late Pleistocene and modern proboscideans, based on genetic data: † Mammut americanum (American mastodon) † Notiomastodon platensis (South American gomphothere) † Mammuthus columbi (Columbian mammoth) † Mammuthus primigenius ( woolly mammoth ) Elephas maximus (Asian elephant) † Palaeoloxodon antiquus (European straight-tusked elephant) Loxodonta cyclotis (African forest elephant) Loxodonta africana (African bush elephant) Within six weeks from 2005–2006, three teams of researchers independently assembled mitochondrial genome profiles of

10353-414: 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

10472-588: The most recent survival of any known mammoth population. Wrangel Island is thought to have become separated from the mainland by 12000 BCE. It was assumed that Wrangel Island mammoths ranged from 180–230 cm (71–91 in) in shoulder height and were for a time considered "dwarf mammoths". However this classification has been re-evaluated and since the Second International Mammoth Conference in 1999, these mammoths are no longer considered to be true "dwarf mammoths", as their size falls within

10591-411: The overhairs, bicoloured, nonpigmented and mixed red-brown guard hairs, and nonpigmented underhairs, which would give a light overall appearance. 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

10710-541: The range of that of mainland Siberian woolly mammoths. Woolly mammoth 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

10829-406: 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

10948-422: The sensitive and muscular trunk worked as a limb-like organ with many functions. It was used for manipulating objects, and in social interactions. The well-preserved foot of the adult male " Yukagir mammoth " shows that the soles of the feet contained many cracks that would have helped in gripping surfaces during locomotion. Like modern elephants, woolly mammoths walked on their toes and had large, fleshy pads behind

11067-415: 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

11186-439: The shoulder and weighed about 350–400 kilograms (770–880 lb). This species became extinct around 1 million years ago, being replaced by Stegodon florensis. Stegodon florensis shows a progressive size reduction with time, with the earlier Middle Pleistocene subspecies Stegodon florensis florensis estimated to be around 50% the size of mainland Stegodon species with a shoulder height of around 190 cm (6.2 ft) and

11305-548: The shoulders, with a body mass of around 5 tons. Other smaller unnamed Stegodon species are also known from the Early Pleistocene on the island. The extinct dwarf elephant species Stegoloxodon indonesicus is also known from the Early Pleistocene of Java, which is probably closely related to S. celebensis from Sulawesi, but whose relationships to other elephants are obscure. The species Stegodon sumbaensis of an uncertain Middle-Late Pleistocene age from Sumba

11424-468: The size of its mainland ancestor. The remains, though temporally poorly constrained, are suggested to be Late Pleistocene age. Possible tracks produced by these dwarf elephants have been reported from the southwest of the island. On Tilos , the species Palaeoloxodon tiliensis has been described from remains found in Charkadio cave. This species was medium-sized, around 10% the size of P. antiquus , with

11543-422: The skull, and less torque would occur than with straight tusks. The tusks were usually asymmetrical and showed considerable variation, with some tusks curving down instead of outwards and some being shorter due to breakage. Calves developed small milk tusks a few centimetres long at six months old, which were replaced by permanent tusks a year later. Tusk growth continued throughout life, but became slower as

11662-445: 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. The completion of

11781-410: The species has been estimated at 2.8–3.15 m (9 ft 2 in – 10 ft 4 in) with a weight of 4.5–6 t (9,900–13,200 lb), with females being smaller like living elephants, with a shoulder height of 2.3–2.6 m (7 ft 7 in – 8 ft 6 in) and a weight of 2.8–4 t (6,200–8,800 lb). This size is comparable to the largest living elephant species,

11900-401: The stomach and faecal matter in the intestines of the mammoth calf " Lyuba ". The faecal matter may have been eaten by "Lyuba" to promote development of the intestinal microbes necessary for digestion of vegetation, as is the case in modern elephants. An isotope analysis of woolly mammoths from Yukon showed that the young nursed for at least 3 years and were weaned and gradually changed to

12019-508: The thermal balance of the animal. 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. 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

12138-429: The time of arrival of modern humans to the islands. In Indonesia and the Philippines, evidence of a succession of distinct endemic island faunas has been found, including dwarfed elephants and species of Stegodon . During the late Early Pleistocene, Flores was inhabited by the dwarf species Stegodon sondaarii , around 15% of the size of mainland Stegodon species, which was around 120 cm (3.9 ft) tall at

12257-415: 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 (30 in) long, whereas the trunk of the adult "Liakhov mammoth" was 2 m (6 ft 7 in) long. The well-preserved trunk of a juvenile specimen nicknamed " Yuka " was described in 2015, and it was shown to possess

12376-841: The toes. Like modern elephants, woolly mammoths were likely very social and lived in matriarchal (female-led) family groups. This is supported by fossil assemblages and cave paintings showing groups, implying that most of their other social behaviours were likely similar to those of modern elephants. How many mammoths lived at one location at a time is unknown, as fossil deposits are often accumulations of individuals that died over long periods of time. The numbers likely varied by season and lifecycle events. Modern elephants can form large herds, sometimes consisting of multiple family groups, and these herds can include thousands of animals migrating together. Mammoths may have formed large herds more often, since animals that live in open areas are more likely to do this than those in forested areas. Trackways made by

12495-419: The tooth closely resembling those of the species P. creutzburgi from Crete (which is adjacent to Kasos) in size and shape, it has been referred to as P. aff. creutzburgi. The Cyprus dwarf elephant ( Palaeoloxodon cypriotes ) survived at least until 12,000 years ago, around the time of arrival of modern humans to Cyprus (who may have hunted it), making it one of the latest surviving dwarf elephants. It

12614-440: 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. 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

12733-711: The weight of the head. The short and tall skulls of woolly and Columbian mammoths ( Mammuthus columbi ) were the culmination of this process. The first known members of the genus Mammuthus are the African species Mammuthus subplanifrons from the Pliocene , 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

12852-463: The wool's insulation, repelled water, and given the fur a glossy sheen. Preserved woolly mammoth fur is orange-brown, but this is believed to be an artefact from the bleaching of pigment during burial. The amount of pigmentation varied from hair to hair and within each hair. A 2006 study sequenced the Mc1r gene (which influences hair colour in mammals) from woolly mammoth bones. Two alleles were found:

12971-493: The woolly mammoth from ancient DNA , which allowed them to confirm the close evolutionary relationship between mammoths and Asian elephants ( Elephas maximus ). A 2015 DNA review confirmed Asian elephants as the closest living relative of the woolly mammoth. African elephants ( Loxodonta africana ) branched away from this clade around 6 million years ago, close to the time of the similar split between chimpanzees and humans. A 2010 study confirmed these relationships and suggested

13090-448: 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. By the early 20th century, the taxonomy of extinct elephants was complex. In 1942, American palaeontologist Henry Fairfield Osborn 's posthumous monograph on

13209-403: Was dismissed in a 2012 study. 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"

13328-885: Was identified as an extinct species of elephant by Georges Cuvier in 1796. 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 . The woolly mammoth was roughly the same size as modern African elephants . Males reached shoulder heights between 2.67 and 3.49 m (8 ft 9 in and 11 ft 5 in) and weighed between 3.9 and 8.2 t (3.8 and 8.1 long tons; 4.3 and 9.0 short tons). Females reached 2.3–2.6 m (7 ft 7 in – 8 ft 6 in) in shoulder heights and weighed between 2.8–4 t (2.8–3.9 long tons; 3.1–4.4 short tons). A newborn calf weighed about 90 kg (200 lb). The woolly mammoth

13447-412: Was not widely accepted at the time. Following Cuvier's identification, the 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

13566-477: Was only modestly dwarfed compared to its large continental ancestor, having a reconstructed shoulder height of 2.4–2.8 metres (7.9–9.2 ft), for males and around 2 metres (6.6 ft) for females. During the Holocene , woolly mammoths ( Mammuthus primigenius ) lived on Wrangel Island in the Arctic Ocean , surviving thousands of years after the extinction of mainland woolly mammoths until around 2000 BCE,

13685-521: Was present at some point in the Pleistocene. It has an estimated body mass of around 400 kilograms (880 lb). Some species of the stegodontid Stegolophodon from the Middle Miocene of Japan around 16 million years ago have been suggested to exhibit insular dwarfism, appearing to exhibit size reduction over time, which would make them the oldest known proboscideans to do so. During Pliocene-Early Pleistocene (from around 4-1 million years ago),

13804-456: Was similar to that of a modern elephant, and that it was born in spring. The best-preserved head of a frozen adult specimen, that of a male nicknamed the "Yukagir mammoth", shows that woolly mammoths had temporal glands between the ear and the eye. This feature indicates that, like bull elephants, male woolly mammoths entered " musth ", a period of heightened aggressiveness. The glands are used especially by males to produce an oily substance with

13923-476: Was subsequently used. 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. Where and how the word "mammoth" originated is unclear. According to the Oxford English Dictionary , it comes from an old Vogul word mēmoŋt , "earth-horn". It may be a version of mehemot ,

14042-407: 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

14161-401: Was well adapted to the cold environments present during glacial periods , including 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

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