page_title: Woolly mammoth
section_title: Evolution
hierarchical_section_title: Woolly mammoth / Taxonomy / Evolution
caption_attribution_description: English: Figure of the jaw of an Indian elephant and the fossil Jaw of a mammoth from Cuvier's 1798–99 paper on living and fossil elephants Magyar: Cuvier illusztrációja az indiai elefánt és a mamut álkapcsáról 1799-ből
context_page_description: The woolly mammoth is an extinct species of 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 Asia. Its closest extant relative is the Asian elephant. 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 Alaska, as well as skeletons, teeth, stomach contents, dung, and depiction from life in prehistoric cave paintings. Mammoth remains had long been known in Asia before they became known to Europeans in the 17th century. The origin of these remains was long a matter of debate, and often explained as being remains of legendary creatures. The mammoth was identified as an extinct species of elephant by Georges Cuvier in 1796.
  The woolly mammoth was roughly the same size as modern African elephants. Males reached shoulder heights between 2.7 and 3.4 m and weighed up to 6 metric tons.
context_section_description: The earliest known members of the Proboscidea, the clade which contains modern elephants, existed about 55 million years ago around the Tethys Sea. The closest known relatives of the Proboscidea are the sirenians (dugongs and manatees) and the hyraxes (an order of small, herbivorous mammals). The family Elephantidae existed 6 million years ago in Africa and includes the modern elephants and the mammoths. Among many now extinct clades, the mastodon (Mammut) is only a distant relative of the mammoths, and part of the separate family Mammutidae, which diverged 25 million years before the mammoths evolved. The following cladogram shows the placement of the genus Mammuthus among other proboscideans, based on characteristics of the hyoid bone in the neck:
  In 2005, researchers assembled a complete mitochondrial genome profile of the woolly mammoth, which allowed them to trace 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. Before the publication of the Neanderthal genome, many researchers expected the first fully sequenced nuclear genome of an extinct species would be that of the mammoth. A 2010 study confirmed these relationships, and suggested the mammoth and Asian elephant lineages diverged 5.8–7.8 million years ago, while African elephants diverged from an earlier common ancestor 6.6–8.8 million years ago. In 2008, much of the woolly mammoth's chromosomal DNA was mapped. The analysis showed that the woolly mammoth and the African elephant are 98.55% to 99.40% identical. The team mapped the woolly mammoth's nuclear genome sequence by extracting DNA from the hair follicles of both a 20,000-year-old mammoth retrieved from permafrost and another that died 60,000 years ago. In 2012, proteins were confidently identified for the first time, collected from a 43,000-year-old woolly mammoth.
  Since many remains of each species of mammoth are known from several localities,  reconstructing the evolutionary history of the genus through morphological studies is possible. Mammoth species can be identified from the number of enamel ridges (or lamellar plates) on their molars; primitive species had few ridges, and the number increased gradually as new species evolved to feed on more abrasive food items. The crowns of the teeth became deeper in height and the skulls became taller to accommodate this. At the same time, the skulls became shorter from front to back to minimise the weight of the head. 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 the earlier type, becoming 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. The Columbian mammoth evolved from a population of M. trogontherii that had crossed the Bering Strait and entered North America about 1.5 million years ago; it retained a similar number of molar ridges. Mammoths derived from M. trogontherii evolved molars with 26 ridges 400,000 years ago in Siberia and became the woolly mammoth. Woolly mammoths entered North America about 100,000 years ago. Asian elephants have a variable number of enamel ridges that is intermediate between those of the Columbian/steppe mammoth and the wooll
caption: Georges Cuvier's 1796 comparison between the mandible of a woolly mammoth (bottom left and top right) and an Indian elephant (top left and bottom right)
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