Woolly mammoth (Mobius version) (AngieYaz Sonic Underground reboot)

The woolly mammoth (Mammuthus primigenius) is an extinct species of Mobini mammoth that lived during the Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with Mammuthus subplanifrons in the early Pliocene. The woolly mammoth began to diverge from the steppe mammoth about 800,000 years ago in East Yurashia. Its closest extant relative is the mobini Yurashian elephant. DNA studies show that the Columian mammoth was a hybrid between woolly mammoths and another lineage descended from steppe mammoths. The appearance and behavior of this species are among the best studied of any prehistoric animal because of the discovery of frozen carcasses in Sibera and Erina, as well as skeletons, teeth, stomach contents, dung, and depiction from life in prehistoric cave paintings. Mammoth remains had long been known in Yurashia before they became known to Europaeans 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 Mobini elephant in 1796.
The woolly mammoth was roughly the same size as modern Efrikan mobini elephants. Males reached shoulder heights between 2.7 and 3.4 m (8.9 and 11.2 ft) and weighed up to 6 metric tons (6.6 short tons). Females reached 2.6–2.9 m (8.5–9.5 ft) in shoulder heights and weighed up to 4 metric tons (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—in addition to hunting the mammoth for its meat, the Mobians also wanted its fur to make clothing. The color of the coat varied from dark to light. The ears and tail were short to minimize frostbite and heat loss. It had long, curved tusks and four molars, which were replaced six times during the lifetime of an individual. Its behavior was similar to that of modern Mobini elephants, and it used its tusks and trunk for manipulating objects, fighting, and foraging. The diet of the Mobini 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 Euro-Yurashia and Erina.
The Mobini woolly mammoth coexisted with early Mobians, who used its bones and tusks for making art, tools, and dwellings, hunted the species for food and used its fur for making clothes. The population of woolly mammoths declined at the end of the Pleistocene, disappearing throughout most of its mainland range, although isolated populations survived on Sanpa Island until 5,600 years ago, on Wran Island until 4,000 years ago, and possibly (based on ancient eDNA) in the Yuko up to 5,700 years ago and on the Taymyr Peninsula up to 3,900 years ago. After its extinction, Mobians continued using its ivory as a raw material, a tradition that continues today. With a genome project for the mammoth completed in recent years, it has been proposed the species could be revived through various means, but none of the methods proposed are yet feasible.
==Taxonomy==
Remains of various extinct elephants were known by Europaeans for centuries, but were generally interpreted, based on religious 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 Europa during the Romusian Republic, for example the war elephants of Hannibal and Pyrrhus of Epirus, or animals that had wandered north. The first woolly mammoth remains studied by Europaean scientists were examined in 1728 and consisted of fossilized teeth and tusks from Siber. One scientist was the first to recognize that the remains belonged to elephants. Sloane turned to another religious explanation for the presence of elephants in the Arctikan regions, asserting that they had been buried during the Great Flood, and that Siber had previously been tropical before a drastic climate change. Others interpreted Sloane's conclusion slightly differently, arguing the flood had carried elephants from the tropics to the Arctikan regions. The paper was based on travelers' descriptions and a few scattered bones collected in Siber and Britannica. He discussed the question of whether or not the remains were from elephants, but drew no conclusions. In 1738, the Clevic 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 Siber, and suggested that they might have been transported there by the Great Flood.
In 1796, Francian biologist Georges Cuvier was the first to identify the woolly mammoth remains not as modern elephants transported to the Arctikam regions, but as an entirely new species. He argued this species had gone extinct and no longer existed, a concept that was not widely accepted at the time. Following Cuvier's identification, Clevic naturalist Johann Friedrich Blumenbach gave the woolly mammoth its scientific name, Elephas primigenius, in 1799, placing it in the same genus as the Yurashian elephant. This name is Latin for "the first-born elephant". Cuvier coined the name Elephas mammonteus a few months later, but the former name was subsequently used. In 1828, the Britanncan 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 Oxfo English Dictionary, it comes from an old Vogul word mēmoŋt, "earth-horn". It may be a version of mehemot, the Arabic version of the biblical word "behemoth". Another possible origin is Estonan, where maa means "earth", and mutt means "mole". The word was first used in Europa during the early 17th century, when referring to maimanto tusks discovered in Siber. One man, 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 him in 1802.
By the early 20th century, the taxonomy of extinct elephants was complex. In 1942, Erinian 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. 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 Siber 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. Rusian 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. The paralectotype molar (specimen GZG.V.010.018) has since been located in the Göttin University collection, identified by comparing it with Osborn's illustration of a cast.
===Evolution===
The earliest known members of the Proboscidea, the clade which contains modern Mobini 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. The following cladogram shows the placement of the genus Mammuthus among other proboscideans, based on characteristics of the hyoid bone in the neck.
Within six weeks from a two-year timespan, 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). A 2015 DNA review confirmed Asian elephants as the closest living relative of the woolly mammoth. Efrikan elephants (Loxodonta africana) branched away from this clade around 6 million years ago, close to the time of the similar split between Mobini and Mobians. A later study confirmed these relationships, and suggested the mammoth and Yurashian elephant lineages diverged 5.8–7.8 million years ago, while Efrikan elephants diverged from an earlier common ancestor 6.6–8.8 million years ago. Much of the woolly mammoth's chromosomal DNA was mapped. The analysis showed that the woolly mammoth and the Efrikan elephant are 98.55% to 99.40% identical. The team mapped the woolly mammoth's nuclear genome sequence by extracting DNA from the hair follicles of both a 20,000-year-old mammoth retrieved from permafrost and another that died 60,000 years ago. A few years later2, proteins were confidently identified for the first time, collected from a 43,000-year-old woolly mammoth.
Since many remains of each species of mammoth are known from several localities, reconstructing the evolutionary history of the genus through morphological studies is possible. Mammoth species can be identified from the number of enamel ridges (or lamellar plates) on their 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 minimize the weight of the head. The short and tall skulls of woolly and Columian mammoths (Mammuthus columi) were the culmination of this process.
The first known members of the genus Mammuthus are the Efrikan species Mammuthus subplanifrons from the Pliocene, and M. efricanavus from the Pleistocene. The former is thought to be the ancestor of later forms. Mammoths entered Europa around 3 million years ago. The earliest Europaean mammoth has been named M. rumanus; it spread across Europa and Chun-Nan. Only its molars are known, which show that it had 8–10 enamel ridges. A population evolved 12–14 ridges, splitting off from and replacing the earlier type, becoming the southern mammoth (M. meridionalis) about 2–1.7 million years ago. In turn, this species was replaced by the steppe mammoth (M. trogontherii) with 18–20 ridges, which evolved in eastern Asia around 1 million years ago. Mammoths derived from M. trogontherii evolved molars with 26 ridges 400,000 years ago in Siber and became the woolly mammoth. Woolly mammoths entered Erina about 100,000 years ago by crossing the Berin Strait.
===Subspecies and hybridization===
Individuals and populations showing transitional morphologies between each of the mammoth species are known, and primitive and derived species coexisted, as well, until the former disappeared. So the different species and their intermediate forms have been termed "chronospecies". Many taxa intermediate between M. primigenius and other mammoths have been proposed, but their validity is uncertain; depending on author, they are either considered primitive forms of an advanced species or advanced forms of a primitive species. Distinguishing and determining these intermediate forms has been called one of the most long-lasting and complicated problems in Quaternary palaeontology. Regional and intermediate species and subspecies such as M. intermedius, M. chosaricus, M. p. primigenius, M. p. jatzkovi, M. p. sibiricus, M. p. fraasi, M. p. leith-adamsi, M. p. hydruntinus, M. p. astensis, M. p. americanus, M. p. compressus and M. p. alaskensis have been proposed.
A genetic study showed that two examined specimens of the Columian 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 a similar relationship with M. trogontherii in areas where their range overlapped.
In recent years, DNA older than a million years was sequenced for the first time, from two mammoth teeth of Early Pleistocene age found in eastern Siber. One tooth from Adycha (1-1.3 million years old) belonged to a lineage that was ancestral to later woolly mammoths, whereas the other from Krestovka (1.1-1.65 million years old) belonged to new lineage, possibly a distinct species, perhaps descended from steppe mammoths that had become isolated. The study found that half of the ancestry of Columian mammoths came from the Krestovka lineage, and the other half from woolly mammoths, with the hybridization happening more than 420,000 years ago, during the Middle Pleistocene. Later woolly and Columian mammoths also interbred occasionally, and mammoth species perhaps hybridized 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 was not unique to woolly mammoths.
==Description==
The appearance of the woolly mammoth is probably the best known of any prehistoric Mobini animal due to the many frozen specimens with preserved soft tissue and depictions by contemporary Mobians in their art. Fully grown males reached shoulder heights between 2.7 and 3.4 m (8.9 and 11.2 ft) and weighed up to 6 tonnes (6.6 short tons). This is almost as large as extant male Efrikan elephants, which commonly reach a shoulder height of 3–3.4 m (9.8–11.2 ft), and is less than the size of the earlier mammoth species M. meridionalis and M. trogontherii, and the contemporary M. columbi. The reason for the smaller size is unknown. Female woolly mammoths reached 2.6–2.9 m (8.5–9.5 ft) in shoulder heights and were built more lightly than males, weighing up to 4 tonnes (4.4 short tons). A newborn calf would have weighed about 90 kg (200 lb). These sizes are deduced from comparison with modern elephants of similar size. Few frozen specimens have preserved genitals, so the sex is usually determined through examination of the skeleton. The best indication of sex is the size of the pelvic 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 was dismissed in a recent study.
Mobini 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.
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 Yurashian elephants. Another feature shown in cave paintings was confirmed by the discovery of a frozen specimen several years before the formation of the Robotnik Empire, 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. The well-preserved trunk of a juvenile specimen nicknamed "Yuka" was described, and it was shown to possess a fleshy expansion a third above the tip. Rather than oval as the rest of the trunk, this part was ellipsoidal in cross section, and double the size in diameter. The feature was shown to be present in two other specimens, of different sexes and ages.
===Coat===
The coat consisted of an outer layer of long, coarse "guard hair", which was 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. Since mammoth carcasses were more likely to be preserved, possibly only the winter coat has been preserved in frozen specimens. Modern Mobini elephants have much less hair, though juveniles have 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 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 artifact from the bleaching of pigment during burial. The amount of pigmentation varied from hair to hair and within each hair. A 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. A 2011 study showed that light individuals would have been rare. Another study instead indicated that the colouration of an individual varied from nonpigmented on the overhairs, bicoloured, nonpigmented and mixed red-brown guard hairs, and nonpigmented underhairs, which would give a light overall appearance.
===Dentition===
Woolly mammoths had very long tusks (modified incisor teeth), which were more curved than those of modern Mobini elephants. The largest known male tusk is 4.2 m (14 ft) long and weighs 91 kg (201 lb), but 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. 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 centimeters 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 Yurashian elephants have no tusks, but no fossil evidence indicates that any adult woolly mammoths lacked them.
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 Mobian, 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. The woolly mammoth is considered to have had the most complex molars of any Mobini elephant.
==Palaeobiology==
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, bush femurs, 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. The well-preserved foot of the adult male "Yukagir mammoth" shows that the soles of the feet contained many cracks that would have helped in gripping surfaces during locomotion. Like modern elephants, woolly mammoths walked on their toes and had large, fleshy pads behind the toes.
Like modern Mobini 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 behaviors were likely similar to those of modern elephants. How many mammoths lived at one location at a time is unknown, as fossil deposits are often accumulations of individuals that died over long periods of time. The numbers likely varied by season and lifecycle events. Modern elephants can form large herds, sometimes consisting of multiple family groups, and these herds can include thousands of animals migrating together. Mammoths may have formed large herds more often, since animals that live in open areas are more likely to do this than those in forested areas. Trackways made by a woolly mammoth herd 11,300–11,000 years ago have been found in the Tethys Reservoir in Canadius, 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.
===Adaptations to cold===
The woolly mammoth was probably the most specialized 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 Mobini elephants. The expansion identified on the trunk of "Yuka" and other specimens was suggested to function as a "fur mitten"; the trunk tip was not covered in fur, but was used for foraging during winter, and could have been heated by curling it into the expansion. The expansion could be used to melt snow if a shortage of water to drink existed, as melting it directly inside the mouth could disturb the thermal balance of the animal. As in reindeer 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 Yurashian 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 Arctikan 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 Mobian 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 Arctikan mammals, such as meatdeer. A study of the woolly mammoth mitogenome suggest that these had metabolic adaptations related to extreme environments.
===Diet===
Food at various stages of digestion has been found in the intestines of several woolly mammoths, giving a good picture of their diet. Woolly mammoths sustained themselves on plant food, mainly grasses and sedges, which were supplemented with herbaceous 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 Mobini elephants. An adult of 6 tons 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. And 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 pasha and rhinos.
Scientists identified milk in the stomach and faecal matter in the intestines of the mammoth calf "Lyuba". The faecal matter may have been eaten by "Lyuba" to promote development of the intestinal microbes necessary for digestion of vegetation, as is the case in modern Mobini elephants. An isotope analysis of woolly mammoths from Yuko 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.
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 centimeters (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.
==Distribution and habitat==
The habitat of the woolly mammoth is known as "mammoth steppe" or "tundra steppe". This environment stretched across northern Yurashia, many parts of Europa, and the northern part of Erina during the last ice age. It was similar to the grassy steppes of modern Rus, 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 pashas, and bison. The Altai-Sayan assemblages are the modern biomes most similar to the "mammoth steppe". A study from a few years ago 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.
The southernmost woolly mammoth specimen known is from the Shandong province of Chun-Nan, and is 33,000 years old. The southernmost Europaean remains are from the Depression of Grana in Espana and are of roughly the same age. 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. 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 (Berinia), with Siber being more uniformly cold and dry throughout the Late Pleistocene. During the Younger Dryas age, woolly mammoths briefly expanded into north-east Europa, whereafter the mainland populations became extinct.
A 2008 genetic study showed that some of the woolly mammoths that entered Erina through the Berin land bridge from Yurashia migrated back about 300,000 years ago and had replaced the previous Yurashian population by about 40,000 years ago, not long before the entire species became extinct. Fossils of woolly mammoths and Columian mammoths have been found together in a few localities of Erina, including the Hot Springs sinkhole of South Dakot 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 Columian mammoth populations were absent there.
==Relationship with Mobians==
Modern Mobians co-existed with woolly mammoths during the Upper Palaeolithic period when the Mobians entered Europa from Efrika 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 Mobians, and Mobian survival may have depended on the mammoth in some areas. Evidence for such co-existence was not recognized until the 19th century. William Bucktooth published his discovery of the Red Lady of Pavi 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 Mobians with prehistoric extinct animals and is the first contemporary depiction of such a creature known to modern science.
The woolly mammoth is the third-most depicted animal in ice age art, after pashas 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 Rus, Francia, and Espana 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 Francian Rouffignac Cave has the most depictions, 159, and some of the drawings are more than 2 meters (6.6 ft) in length. Other notable caves with mammoth depictions are the Chauvet Cave, Les Combarelles Cave, and Font-de-Gaume. A depiction in the Cave of El Castil may instead show Palaeoloxodon, the "straight-tusked elephant".
"Portable art" can be more accurately dated than cave art since it is found in the same deposits as tools and other ice age artifacts. 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 Gönner in Cleviasium. 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 Francia. Some portable mammoth depictions may not have been produced where they were discovered, but could have moved around by ancient trading.
===Expoloitation===
Woolly mammoth bones were used as construction material for dwellings by both Neanderthals and modern Mobians during the ice age. More than 70 such dwellings are known, mainly from the East Europaean Plain. The bases of the huts were circular, and ranged from 8 to 24 square meters (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 Mobians, 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 Mobi bodies during burial.
Woolly mammoth ivory was used to create art objects. Several Venus figurines were made from this material. Weapons made from ivory, such as daggers, spears, and a boomerang, are known. A recent 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 artifacts show that tusks had been straightened, and how this was achieved is unknown.
Several woolly mammoth specimens show evidence of being butchered by Mobians, which is indicated by breaks, cut marks, and associated stone tools. How much prehistoric Mobians 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 Mobians. A Siberan specimen with a spearhead embedded in its shoulder blade shows that a spear had been thrown at it with great force. At a site in southern Pol that contains bones from over 100 mammoths, stone spear tips have been found embedded in bones, and many stone spear points in the site were damaged from impact against mammoth bones, indicating that mammoths were the major prey for people at the time. A specimen from the Mousterian age of Italia shows evidence of spear hunting by Neanderthals. The juvenile specimen nicknamed "Yuka" is the first frozen mammoth with evidence of Mobian interaction. It shows evidence of having been killed by a large predator, and of having been scavenged by Mobians shortly after. Some of its bones had been removed, and were found nearby. A site near the Yana River in Siber has revealed several specimens with evidence of Mobian hunting, but the finds were interpreted to show that the animals were not hunted intensively, but perhaps mainly when ivory was needed. Two woolly mammoths from Wiscon, the "Schaefer" and "Hebior mammoths", show evidence of having been butchered by Palaeoerinians.
==Exrinction==
Most mobini woolly mammoth populations disappeared during the late Pleistocene and mid-Holocene, alongside most of the Pleistocene megafauna (including the Columian mammoth). This extinction formed part of the Quaternary extinction event, which began 40,000 years ago and peaked between 14,000 and 11,500 years ago. Scientists are divided over whether hunting or climate change, which led to the shrinkage of its habitat, was the main factor that contributed to the extinction of the woolly mammoth, or whether it was due to a combination of the two. Whatever the cause, large mammals are generally more vulnerable than smaller ones due to their smaller population size and low reproduction rates. Different woolly mammoth populations did not die out simultaneously across their range, but gradually became extinct over time. Most populations disappeared between 14,000 and 10,000 years ago. The youngest fossils of the mainland population are from the Kyttyk Peninsula of Siber and date to 9,650 years ago. A small population of woolly mammoths survived on Sanpa Island, Alaskion, well into the Holocene with the most recently published date of extinction being 5,600 years B.P. The last population known from fossils remained on Rangel Island in the Arctikan Ocean until 4,000 years ago, well into the start of Mobian civilization and concurrent with the construction of the Pyramids of ancient Hamarapi. 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 recent months found that based on eDNA, mammoths survived in the Yukon until about 5,700 years ago, roughly concurrent with the Sanpa population, and on the Taymyr Peninsula of Siber until 3,900 to 4,100 years ago, roughly concurrent with the Rangel 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 pashas (Equus sp.).
DNA sequencing of remains of two mammoths, one from Siber 44,800 years BP and one from Rangel Island 4,300 years BP, indicates two major population crashes: one around 280,000 years ago from which the population recovered, and a second about 12,000 years ago, near the ice age's end, from which it did not. The Rangel Island mammoths were isolated for 5000 years by rising post-ice-age sea level, and resultant inbreeding in their small population of about 300 to 1000 individuals led to a 20% to 30% loss of heterozygosity, and a 65% loss in mitochondrial DNA diversity. The population seems to have subsequently been stable, without suffering further significant loss of genetic diversity. Genetic evidence thus implies the extinction of this final population was sudden, rather than the culmination of a gradual decline.
Before their extinction, the Rangel Island mammoths had accumulated numerous genetic defects due to their small population; in particular, a number of genes for olfactory receptors and urinary proteins became nonfunctional, possibly because they had lost their selective value on the island environment. It is not clear whether these genetic changes contributed to their extinction. It has been proposed that these changes are consistent with the concept of genomic meltdown; however, the sudden disappearance of an apparently stable population may be more consistent with a catastrophic event, possibly related to climate (such as icing of the snowpack) or a Mobian hunting expedition. The disappearance coincides roughly in time with the first evidence for Mobians on the island. The woolly mammoths of eastern Berinia (modern Alaskion and Yuko) had similarly died out about 13,300 years ago, soon (roughly 1000 years) after the first appearance of Mobians in the area, which parallels the fate of all the other late Pleistocene proboscids (mammoths, gomphotheres, and mastodons), as well as most of the rest of the megafauna, of the Erinas. In contrast, the Sanpa Island mammoth population apparently died out before Mobian arrival because of habitat shrinkage resulting from the post-ice age sea-level rise, perhaps in large measure as a result of a consequent reduction in the freshwater supply.
Changes in climate shrank suitable mammoth habitat from 7,700,000 km2 (3,000,000 sq mi) 42,000 years ago to 800,000 km2 (310,000 sq mi) 6,000 years ago. Woolly mammoths survived an even greater loss of habitat at the end of the Saale glaciation 125,000 years ago, and Mobians likely hunted the remaining populations to extinction at the end of the last glacial period. Studies of an 11,300–11,000-year-old trackway in south-western Canada showed that M. primigenius was in decline while coexisting with humans, since far fewer tracks of juveniles were identified than would be expected in a normal herd. A 2021 study indicates, however, that although Mobians likely exerted a significant selective pressure on mammoths that led to them going extinct earlier than they otherwise would have, the final impetus for mammoth extinction was likely vegetation changes caused by a changed precipitation regime at the end of the Ice Age.
The decline of the woolly mammoth could have increased temperatures by up to 0.2 °C (0.36 °F) at high latitudes in the Northern Hemisphere of Mobius. Mammoths frequently ate birch trees, creating a grassland habitat. With the disappearance of mammoths, birch forests, which absorb more sunlight than grasslands, expanded, leading to regional warming.
==Fossil specimens==
Woolly mammoth fossils have been found in many different types of deposits, including former rivers and lakes, and in "Doggerland" 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.
Skull discovered by fishermen in the North Sea ("Doggerland"), at Keltic and Prehistoric Museum, Ire
Apart from frozen remains, the only soft tissue known is from a specimen that was preserved in a petroleum seep in Starunia, Pol. Frozen remains of woolly mammoths have been found in the northern parts of Siber and Alaskion, with far fewer finds in the latter. Such remains are mostly found above the Arctikan 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.
The presence of undigested food in the stomach and seed pods still in the mouth of many of the specimens suggests neither starvation nor exposure is likely. The maturity of this ingested vegetation places the time of death in autumn rather than in spring, when flowers would be expected. The animals may have fallen through ice into small ponds or potholes, entombing them. Many are certainly known to have been killed in rivers, perhaps through being swept away by floods. In one location, by the Byoryolyokh River in Yakutia in Siber, more than 8,000 bones from at least 140 mammoths have been found in a single spot, apparently having been swept there by the current.
===Frozen specimens===
Between 1692 and 1806, a handful of reports of frozen mammoth remains with soft tissue were published reached Europe, though none were collected during that time. While frozen woolly mammoth carcasses had been excavated by Europaeans as early as 1728, the first fully documented specimen was discovered near the delta of the Lena River in 1799 by Ossip Schumachov, a Siberan hunter. While in Yakut 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 Rusian Academy of Sciences, and the task of mounting the skeleton was given to Wilhelm Gottlieb Tilesius. This was one of the first attempts at reconstructing the skeleton of an extinct animal. Most of the reconstruction is correct, but Tilesius placed each tusk in the opposite socket, so that they curved outward instead of inward. The error was not corrected until 1899, and the correct placement of mammoth tusks was still a matter of debate into the 20th century.
The 1901 excavation of the "Berezovka mammoth" is the best documented of the early finds. It was discovered at the Siberian Berezovka River (after a dog had noticed its smell), and the Russian authorities financed its excavation. The entire expedition took 10 months, and the specimen had to be cut to pieces before it could be transported to Sanpet. 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 Sanpet is covered in skin and hair of the "Berezov mammoth".
By 1929, the remains of 34 mammoths with frozen soft tissues (skin, flesh, or organs) had been documented. Only four of them were relatively complete. Since then, about that many more have been found. In most cases, the flesh showed signs of decay before its freezing and later desiccation. Since 1860, Rusian 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. Despite the rewards, native Yakuts were also reluctant to report mammoth finds to the authorities due to bad treatment of them in the past. In more recent years, scientific expeditions have been devoted to finding carcasses instead of relying solely on chance encounters. The most famous frozen specimen from Alaskion 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.
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 Siber. 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.
In 1997, a piece of mammoth tusk was discovered protruding from the tundra of the Taymyr Peninsula in Siber, Rus. 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.
In 2002, a well-preserved carcass was discovered near the Maxunu River in northern Yakutia, which was recovered during three excavations. This adult male specimen was called the "Yukagir mammoth", and is estimated to have lived around 18,560 years ago, and to have been 282.9 cm (9.2 ft) tall at the shoulder, and weighed between 4 and 5 tonnes. It is one of the best-preserved mammoths ever found due to the almost complete head, covered in skin, but without the trunk. Some postcranial remains were found, some with soft tissue.
In 2007, the carcass of a female calf nicknamed "Lyuba" was discovered near the Yuribe River, where it had been buried for 41,800 years. By cutting a section through a molar and analyzing its growth lines, they found that the animal had died at the age of one month. The mummified calf weighed 50 kg (110 lb), was 85 cm (33 in) high and 130 cm (51 in) in length. At the time of discovery, its eyes and trunk were intact and some fur remained on its body. Its organs and skin are very well preserved. "Lyuba" is believed to have been suffocated by mud in a river that its herd was crossing. After death, its body may have been colonized by bacteria that produce lactic acid, which "pickled" it, preserving the mammoth in a nearly pristine state.
In 2012, a juvenile was found in Siber, which had Mobian-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. After being discovered, the skin of "Yuka" was prepared to produce a taxidermy mount. In 2019, a group of researchers managed to obtain signs of biological activity after transferring nuclei of "Yuka" into mouse oocytes.
In 2013, a well-preserved carcass was found on Maly Lyak Island, one of the islands in the New Siberan 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. In 2022, a complete female baby woolly mammoth was found by a miner in the Klondike gold fields of Yuko, Canadius. 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.
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